Research Programs Constituting U.S. Participation in the International Biological Program

DOCUMENT RESUME

ED 059 903 SE 013 412

TITLE Research Programs Constituting U.S. Participation in the International Biological Program. INSTITUTION National Academy of Sciences--National Research Council, Washington, D.C. Div. of Biology and Agriculture. REPORT NO R-4 PUB DATE 71 NOTE 121p.

EDRS PRICE 4F-$0.65 HC-$6.58 DESCRIPTOPS *Biology; *Ecology; Environmental Education; Genetics; Interdisciplinary Approach; International Programs; Program Descriptions; Resource Materials; Social Sciences; *Urban Population; Zoology IDENTIFIERS *Human Biology; International Biological Program

ABSTRACT The United States contribution to the International Biological Program, which aims to understand more clearly the interrelationships within ecosystems, is centered on multidisciplinary research programs investigating the biological basis of ecological productivity and human welfare. Integrated research programs have been established for the analysis of ecoystems; the study of biological productivity in coastalupwelling ecosystems; the provision of data needed for marine mammal management; the comparison of different ecosystems with similar physical features; the population genetics of South American Indians; the biology of high altitude human populations; and the study of circumpolar peoples. Co-ordirlted research projects have been established on the ecology of nitrogen; the biological control of insect pests; aerobiology; phenology; ecosystem conservation; the conservation of plant genetic materials; the biosocial adaptation of migrant and urban human populations; human nutritional adaptation; and human biological rhythms. For each program there are statements of the research objectives, a summary of progress, a list of publications, and the names of participants. (m) U.S. DEPARTMENT OF HEALTH EDUCATION & WELFARE

_ OFFICE OF EDUCATION THIS DOCUMENT HAS BEEN REPRO DUCED EXACTLY AS RECEIVED FROM THE PERSON OR OF NI2ATION ORIG INATING IT. POIN1- VIEW OR OPIN IONS STATED DO No NECESSARILY REPRESENT OFFICIAL OFFICE OF EDU CATION POSITION OR POLICY

Alit'PROG RV1 ATI HE TER Tif GICAL PR

REPORT NO. 4 OF THE U.S. NATIONP L COMMITTEE FOR THE INTERNATIONAL BIOLOGICAL PROGRAM

U.S. National Committee for the International Biological Program

Division of Biology and Agriculture

National Research Council

NATIONAL ACADEMY OF SCIENCES

Washington, D.C.1971 PRERACE

This report summarizes progress in organizing the research programs that constitute U.S. participation in the International Biological Program and describes the status of research within these multidisciplinary studies. It follows a report entitled Research Studies Constituting the U.S. Contribution to the International Biological Program, which was published in two parts. It differs from the previous report in that it is devoted exclusively to the research programs. The earlier reports included projects judged to be "IBP-related"; files and indices of these are maintained in the office of the U.S. National Committee. International coordination is accomplished through the Program Directors and through the Executive Committee of the U.S. National Committee for the IBP. Those interested in the work of a given program are encouraged to communicate with the appropriate Director. Support for the research is provided directly to the individual programs by a number of federal agencies: National Science Foundation, Department of Agricultuf,..n, Atomic Energy Commission, Smithsonian Institution, Department of the Interior, Depai talent of Com- merce, Department of Defense, Department of Health, Education, and Wellare, and Na- tional Aeronautics and Space Administration. Several private foundations and industrial organizations provide support for components of the programs. The Executive Committee of the U.S. National Committee for the IBP acknowledges with gratitude the efforts of the Program Directors and their colleagues and of the scientists in other countries who have contributed in many ways to the development of the research effort.

Executive Committee of the U.S. National Committee for the IBP Contents

INTRODUCTION INTEGRATED RESEARCH PROGRAMS 3 Environmental Component Analysis of Ecosystems 5 Biological Productivity in Upwelling Ecosystems 37 Marine Mammals 44 Origin and Structure of Ecosystems 46 Human Adaptability Component Population Genetics of South American Indians 53 Biology of Human Populations at High Altitudes 60 International Study of Circumpolar Peoples 65

COORDINATED RESEARCH PROGRAMS 71 Environmental Component Biology and Ecology of Nitrogen 73 Biological Control of Insect Pests 75 Ae robiology 78 Phenology 83 Conservation of Plant Genetic Materials 89 Conservation of Ecosystems 92 Human Adaptability Component Biosocial Adaptation of Migrant and Urban Populations 94 Nutritional Adaptation to the Environment 98 Chronobiology 114

APPENDIXES 117 A Organization of the IBP in the United States 119 B Officers and Members of the Special Committee for the International Biological Program 120 C Countries Participating in the IBP, with Chairmen of National Committees 121 Introduction

During the discussions from which the International cidate the biological basis of productivity and human Biological Program emerged, it was recognized that mul- welfareis zentered on the multidisciplinary research tidisciplinary approaches appropriate to the IBP should programs. Two types of research programs have been or- be encouraged and that these studies would almost cer- ganized: integrated research programs and coordinated tainly continue beyond the formal life of the IBP. research programs. The overall goal is to understand more clearly the inter- Each of the integrated research programs has: relationships within and among ecosystems by: Demonstrated that it is necessary to integrate a Advancing knowledge of the genetic adaptations of number of research projects if the scientific goals of the man. program are to be realized. Advancing knowledge of the physiological and be- Posed questions of appreciable scientific signifi- havorial adaptations of man. cance, thus giving coherence and direction to the suite of Formulating a basis for better understanding the research projects in the program. interactions of components of representative biological Devised a system for coordination and rapid ex- systems. change of information and data so that as work proceeds Exploiting the urderstanding of biological system each participant will profit from the efforts of others. to the end of increasing biological productivity. Provided a means of synthesizing the results of in- Providing bases for predicting the consequences of cluded projects, thus enhancing the total achievement. environmental stresses, whether man-made or natural. Coordinated research programs are an association of Enhancing our ability to better manage our natural related research that when taken collectively make a con- resources. tribution to the field. The emphasis is on coordination of The U.S. response to the challenge of the I BPto elu- closely related, usually ongoing, research. II MI 'DO IFI IIE /OIL Ft 11-11 IF° FRC, IFt WI ENVIRONMENTAL COMPONENT Analysis of Ecosystems

The analysis-of-ecosystems program is designed to pro- reasonably satisfactory so long as the goals of manage- vide basic informatio:i on ecological systems through a ment are centered on single species or populations. comparative study of six major habitat types (biomes). The multiple use of natural renewable resources is in- This knowledge will be useful for solving problems of creasing steadily, however, often combining timber pro- biological production, resource management, and envi- duction, hunting, fishing, and recreation; management ronmental quality. programs attempt to maximize these uses simultaneously. Past studies have analyzed a number of aspects of eco- The contribution of the analysis-of-ecosystems program systems, including kinds of organisms present, their abun- to resource management rests in the manipulation dance, their arrangement in soce, and their food; flow of planned in ecological systems ;--id the production data energy through the system; and cycling of nutrients. Al- that will result. though the data describe what happens, they do not ex- The quality of the environment is another facet of plain why it happenswhy so many species live together, this program. Some types of environmental deterioration why so much plant production is not eaten by animals, are evident to the most casual observer; others of equal why mortality rates are so high among most herbivores, Dr even greater importance are less visible and exert their or why systems persist year after year although individual ?.ffects over decades. As examples of this, one !night list population levels fluctuate between extremes. Finding the thinning of the soil, the decreasisig quality of water the explanations requires more than analysis. It requires supplies, the increasing number of pest species, the dense a synthesis of information into working models describ- algal blooms in lakes, the decreasing number of game fish, ing the structure and functions of whole ecosystems. and other changes in the number and abundance of na- We know that green plants, herbivores, predators, and tive species. decomposers are all interconnected and that production Many problems of environmental deterioration cannot at one level may be greatly influenced by activity in other be understood without a profound knowledge of ecolog- parts of the system. However, little is known about the ical systems. The relationship between cause and effect processes. Studies with mathematical models of systems of deterioration in a system is often difficult to establish. suggest that interrelations may be much more pervasive Usually the cause lies in one part of the system and the than they are now believed to be. response is seen in another. Finding the relationship Strong emphasis is being placed on the physical and would be greatly facilitated by a complete understanding chemical factors of the environment since the ecosystem of ecosystem processes and their couplings. is the combined result of interactions among these factors and biological factors. Such fields as botany, zoology, microbiology, soil science, geology, hydrology, OBJECTIVES meteorology, and limnology are being brought together. Equally important to the combined efforts are several The stated goals of the analysis-of-ecosystems program fields of engineering, particularly systems engineering. are: Understanding the functioning of the ecosystems will To establish a scientific base for programs to main- provide a more valid basis for management of renewable tain or improve environmental quality resources. All management programs for renewabIe re- To derive broad principles of ecosystem structure sources contain ecological concepts that attempt to de- and function through an integration of the results of six scribe the response of the managed population to its biome studies management. At present, the ecological theory that To relate these principles to characteristics of eco- serves this function relates to individuals or to popula- systems (e. g., persistence, stability, maturity, and diver- tions, not to systems as wholes. This limited approach is sity)

5 To develop and refine a generalized adaptable simu- ulation models. The goal is to produce functional models lation model suitable for use in planning studies fornew that serve for the computer simulation of eCosystem development projects processes. In systems analysis, an ecosystem is regarded Each of the biome studies (grasslands, deciduous for- as a number of subsystem,. Since a subsystem is simpler ests, coniferous forests, desert, tundra, and tropical for- than the system as a whole, this smaller unit is moresus- ests) includes among its objectives: ceptible to analytical treatment. A submodel of each To relate productivity, nutrient cycling, energy subsystem can be built, and the submodels can then be flow, and other characteristics of ecosystems in a set of synthesized into a molel of th; vhole. In ecology, the distinct environments complexities of the whole system aie far greater than To determine the driving forces, the processes caus- those with which most other scientists have had to deal, ing transfers of matter and energy among components, and the methods of classical mathematics that physical the nonconcentration charactr dstics, and the controlling scientists can use to combine separate studies into a variables in cach biorne whole have not been applicable. The general availability To determine the ecosystem response to natural of electronic computers has made the systems analysis and man-induced stresses apr ropriate to each biorne approach in ecology a practical possibility. (e.g., large herbivores in the grasslands Uome, extreme Two types of field studies are required: process weather patterns in the desert biome, periodic fluctua- studies and validation studies. The purpose of process tions of rodent populations in the tundra biome, commer- studies is to describe the rate of change of eachecosys- cial use of timber in the coniferous forests biorne, urban- tem component in terms of the present state of thiscom- ization in the deciduous forests biome, and nutrient ponent and all other components of the ecosystem. re:.ention in the wet tropical forests biome) Results of the process studies are used to construct the To understand the land-water interactions charac- ecosystem model. To understand processes, we must do teristic of each biome (e. g., prairie ponds and reservoirs more than measure them. If a single system were analyzed in the grasslancis biome, the abundance of shallow waters completely, there would not be any informationon fac- in the tundra biome, springs and temporary waters in the tors that produced the observed result; and there would desert biome, river systems with anadromous fish popula- not be any means of predicting what would happen if tions in the coniferous forests biome, pollution andeu- system components were altered. Developing this addi- trophication in the deciduous forests biome, and large tional information requires that processes be observedas river systems in the tropical forests biome) they change in response to manipulation of causal fac- To synthesize the results of atese and previous tors and that both the causal factors and the process out- studies into predictive models showing temporal andspa- put be monitored. Various system manipulations are in- tial variation, effects of pollutants and of exploitation, cluded in each of the biome programs. stability, and other ecosystem characteristicsnecessary Since the ecosystem components are toonumerous to for resource management in each biome be studied individually, many of them must be combined for study. This procedure may lead to inaccurate models. Hence, the models must be checked against actual field RESEARCH PLANS conditions. Studies that gather data to check theecosys- tem models are called validation studies. The value of the ecosystem models, developed by the The analysis-of-ecosystems program consists ofa cen- biome studies, depends on how widely such modelscan tral program and studies of ecosystems in six biomes. be applied. It is hoped that one model, with different Most Americans live in and extract the major portion of sets of components and parameters, will apply to all vari- their food and fiber from three of these biomes:grass- ations of a Oven biome. The permissible range of general- lands, deciduous forests, and coniferous forests. These ization for the simulation model will be determined by biomes extend over in the middle range of temperature the validati-)n studies against which it has been checked. and precipitation; the others (tundra, desert, and tropical If the validation sites are a representative sample of the forests) were selected because theyencompass extreme biome and if the simulation model has been satisfactorily environmental conditions. validated against them, then the model'srange of applica- The central program develops the ecosystem studies, bility can be regarded as covering the biomeas a whole. ensures comparability of studies, and provides the synthesis of results needed to develop ecologizal principles based on compaiisons of results from the six biomes. RESEARCH METHODS AND PROGRESS The ecosystem studies makr: extensive use of systems analysis, which includes the orderly and logical organiza- The progress achieved in the central program is best tion of data and development of mathematical and sim- measured by the progress achieved in the biome studies

6 described in the sections that follow. Because funds were tivities having to do with aquatic microbiology, meteor- obtained at different times, the studies are in different ology, modeling, and primary production. stages of development, and synthesis of results has not yet been possible. Although the studies are in different stages, efforts GRASSLANDS BIOME are being made to coordinate them: Subjects related to all six studies are discussed at interbiome meetings, which are attended by specialists The grasslands biome in the United States includes an- from various fields. Wtters discussed include compara- nual, desert, shortgrass, mixed prairie, tallgrass, Palouse, bility of methods, new techniques, and the kinds of arid mountain grasslands. The si;ategy adopted for in- studies needed in the context of the total program. vestigating the physical and functional variations within The need to expedite communication has been rec- this biome called for two kinds of studies: (1) Detailed ognized. First, investigators must be kept informed about process studies are conducted at a single "intensive site." progress of the studies. This need is met by arranging fre- (2) Other process studies and validation studies are con- quent meetings for investigators. Second, research results ducted in a network of sites. must be disseminated rapidly. An interbiome committee The intensive site, known as the Pawnee Site, is in has made significant progress in developing a system for north-central Colorado (Figur:, 1). Field investigations storing and retrieving information. began there in the middle of 1968. The comprehensive Interbiome groups are engaged in coordinating ac- network sites are located throughout the grasslands (Figures 2 and 3). Field investigations began there early in 1970. The general characteristics of these sites are

WYOMING i given in Table 1. Detailed descriptions of the sites are NEBRASKA found in Technical Reports 1 and 36-45 (see Appen- dix A, p. 14).

COLORADO N. Our broad research ohjeLtives are to study various nat- I 'KANSAS ural and man-stressed grassland ecosystems (1) to determine the interrelationships of structure and function, (2) 0-KTATIOMA

NEW MEXICO TEXAS I

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FIGURE 1Pawnee Site. H = heavy herbivore treatment; FIGURE 2 Location of comprehensive network sites in L = light herbivore treatment; M = moderate herbivore relation to the Pawnee Site (Colorado), the Matador Site treatment; S = summer herbivore treatment; W = winter (Saskatchewan, Canada), and the Campana Site (Chihua- herbivore treatment; PVT = private land for aquatic and hua, Mexico). *= first order (intensive);= second order; telemetry study; X = microwatershed; X in a square = en- = third order; 0 = proposed; 0= potential; *= other closed microwatershed. North American grassland IBP sites.

7 Compare: We observe and measure the responses of the system to 7 Annual grassland Ta Boreal Pr. Me different sets of stresses. From detailed analyses of the e' results of the experiments we derive information about HOP LAND OSAGE the effects of driving forces, components, and processes on one another. Further, we examine how these are re- to lated to structure and to community or social interactions. 75 We have designed a variety of initial models of total grassland systems or of their components to provide a focus for our research efforts. These models are mathe- BRIDGER matical expressions of various degrees of complexity and HAYS 45. resolution, and many include probabilistic measurements.

50 PANTEX Much of our analysis effort has been devoted to developing an overall scheme in which to frame our individual

DICKINSON models. Eventually, we translate the mathematical ex-

COTTONWOOD pressions into computer languages to permit simulation

BISON and optimization analyses to be made. From such analy- 14f PAWNEE ses we expect to obtain a basis for redesigning our ex- 25 \4C ALE.' 41 Compare: periments. Mixed prairie JORNADA Palouse Prairie At the Pawnee Site, a shortgrass prairie, we have established a large interdisciplinary team to measure as many as possible of the compartments, driving forces, parameters, and control variables in a set of stressed grasslands.

APRIL MAY JUNE The nature of the research determines the size of the SITE PHENOLOGY leverage date of lilac bloom! areas studied. Examples include (1) detailed soil moisture FIGURE 3Environmental gradients of comprehensive dynamics research in a weighing lysirneter 3 m in diame- network sites. ter, (2) eight 0.5-hectare microwatersheds, (3) 14 1- hectare stress treatment plots, (4) four 10-hectare pas- to determine the variability and magnitude of rates of en . tures used for large-mammal comparative-diet studies, ergy flow an d nutrient cycling, and (3) to encompass (5) three 128-hectare pastures used for domestic-animal these parameters and variables in an overall systems grazing studies, and (6) areas of over 1,000 hectares for model. We have used a systems-analysis approach in our studies involving widely ranging animals (e.g., jackrabbits research, systematically isolating and examining ecosys- and coyotes), and many birds. Under memoranda of tems and their components (see Technical Report 32). agreement with the U.S. Department of Agriculture, ad- We ask the following general questions: ditional areas for research are available on the 6,000- 1. What are the driving forces that make grassland eco- hectare Central Plains Experimental Range and the systems operate? 44,000-hectare western part of the Pawnee National 2. What are the major and minor components of grass- Grassland (see Figure 1 and Technical Report 8). land ecosystems, and what are the changes in magnitude The major stressing or perturbation treatment is that of these components over time? of herbivory, imposed by more than 30 years of known 3. What are the important families or groups of pro- zero, light, moderate, or heavy arazing.by domestic ani- cesses that cause the interaction or linking of the compo- mals. Moisture and nutrient stresses are also imposed on nents of the ecosystem to one another? selected plots. We plan to impose additional stresses, We must also know the changes in certain parameters such as replacement or elimination of different biotic of ecosystems over time, and we must know how plant, components of the community. animal, and microbial community or social interactions In the comprehensive network sites, including all affect processes. Furthermore, we include manipulations major types of United States grasslands, we assemble or treatments by man to determine their influence on smaller interdisciplinary teams to focus primarily on two processes and components. treatments: ungrazed by domestic animals and heavily grazed by domestic animals. Thus, the comprehensive network program includes a subset of treatments found in the Pawnee Site studies. Measuremcnts are made in RESEARCH PLANS the network according to a plan developed and agreed upon by the research participants (see Technical Re- The research objectives, which are broad and diverse, port 35). made it necessary to design experiments with great care. In both the Pawnee Site and the comprehensive net-

10 TABLE 1Characteristics of Comprehensive Network Sites and Intensive Site Annual Elevation Major Grassland Phenology ppt Site (m) Vegetation Type (lilac bloom) (cm) Soil

Comprehensive network sites Arid lands 520 Agropyron spicatum Palouse May 1 30 glaciofluvial ecology site Stipa cornata and aeolian Bison 980 Festuca scabrella Palouse May 20 33 Chernozem Festuca idahoensis Bridger 2,320 Festuca idahoensis Mountain July 1 61 Chernozem Agropyron subsecundum Cottonwood 850 Agropyron smithii Mixed May 20 38 silty clay Stipa viridula Dickinson 850 Stipa comata Mixed May 30 41 Flasher loamy Bouteloua gracilis fine sand Hays 610 Andropogon scoparius Mixed April 25 58 Armo, immature, Andropogon gerardi rocky Hop land 300 Avena fatua Annual April 5 94 Prairie Hordeum jubatum Jornada 1,340 Bouteloua eriopoda Desert April 1 23 alluvial fill Sporobolus flexuosus Osage 380 Andropogon gerardi 1 allgrass April 20 94 Labette, silty Panicum virgatum clay Pantex 1,090 Bouteloua gracilis Shortgrass April 15 53 Pullman, silty Aristida longiseta clay loam Intensive site 1,430 Bouteloua gracilis Shortgrass May 10 30 alluvial plains, Buchloe dactyloides sandy clay loarns

work sites, teams measure the abioti:: components of the 2. Secretarial assistance system and the biotic factors. The difference is primarily 3. Bookkeeping assistance one of extent, the effort being more intense on the 4. Typing assistanc?, Pawnee SRe. Analysis of the results of the Pawnee Site B.Services (George M. Van Dyne) studies provides us with forms of equations in simulation 1. Data processing and analysis (Lewis J. Bledsoe and and optimization models and means to estimate coeffi- Francis Swartzman) cients in those equations. We can construct similar but 2. Sample processing and analysis (Charles L. Streeter) simplified models for the comprehensive network site C.Scientific coordination and information (George M. studies. We can then compare results obtained from run- Var Dyne) ning these different models, test them against the varia- 1. Scientific coordination (Dennis H. Knight) tions of physical and climatic characteristics of the dif- 2. Information dissemination (R. Gerald Wright and ferent grassland sites, and examine differences in their Peter T. Haug) biotic components. This will lead to ecological generaliza- 3. Meetings (R. Gerald Wright and Peter T. Haug) tions about grasslands that we expect will be valuable to D.Systems analysis (George M. Van Dyne) natural resource management. 1. Grasslands seminar (Herbert G. Fisser) The Natural Resource Ecology Laboratory at Colorado 2. Mathematical modeling (George M. Van Dyne, State University, Fort Collins, manages and coordinates Donald A. Jameson, Norman R. French, Gordon the research. George M. Van Dyne is Grasslands Biome Swartzman, Robert C. Francis, Lewis J. Bledsoe, Director. The organizational structure is shown in the and subject-matter specialists) following list. Activities at the first level (Administration, E.Pawnee Site project (Donald A. Jameson) Services, etc.) are phases; each has a director. Those at 1. Facilities, equipment, reference collections the second level are areas; each has a manager. (Robert E. Bement, Freeman M. Smith, and William M. Klein) A. Administration (R. Gerald Wright and Peter T. 7,5a1,g) 2. Remote sensing (Lee D. Miller) 1. Administrative assistance 3. Abiotic-atmospheric (Robert D. Burrnan)

9 4. Abiotic-edaphic (John 0. Reuss) characterize the physical environment. Analysis of histor- 5. Producers (William H. Moir) ical weather data, including temperature, precipitation, 6. Consumers (Kirvin L. Knox) and wind, has suggested that temperature has a strong 7. Decomposers (Kenneth G. Doxtader) seasonal component and lags behind total daily radiation 8. Aquatic (James W. Lavelle) about 1 month. This lag is directly related to the number F. Comprehensive program (Norman R. French) of cloudy days in June that have the effect of lowering 1. Network Studies (Norman R. French) the average daily temperature for that month. For simu- 2. Process Studies (Norman R. French) lation purposes, temperatures can be represented as de- G. International cooperation (George M. Van Dyne) partures from the season component, and the departures can be associated with rainfall periods. The central biome office does some purchasing; pro- Precip7 ation analysis has revealed that regional varia- vides budgeting, administrativc, and chemical analysis tion is greatest from west to east. This agrees with earlier services; and supervises scientific coordination and infor- climatic studies that concluded that, climatically, the mation, preparation of manuscripts and reports, data grasslands are more uniform from south to north. The processing, and systems analysis (see Technical Reports generation of precipitation data for simulation can be rep- 24 and 34). The Pawnee Site operates a field dormitory, resented as a simple first-order Markov process of either kitchen, dining hall, laboratory, and sample processing a rain or a no-rain day. If a day is rainy, the amount of facility. rain is selected from a frequency distribution for that Research investigations are grouped according to driv- day of the year. ing forces and trophic levels. The driving forces include Wind studies show that wind travel contains a seasonal the abiotic components of the environment. The trophic component but is independent of both temperature and levels are producers, consumers, and decomposers. Scien- precipitation. tific coordinators, subject-matter specialists, and interdis- Measurements of overland flow from the microwater- ciplinary specialists help maintain communication and sheds have revealed that it occurs more frequently than coordination among these research groups. one might think, but only locally, reinfiltrating before A list of participants, which includes addresses aild reaching local depressions. areas of specialization, is given in Appendix C (p. 18). In the primary producer studies it has been learned (1) (See Technical Report 24 for detailed vitae and descrip- that shortgrass species respond to significant rainfall tions of work.) Participants contract with the Biome events in less than 72 hours, (2) that heavy rainfall rates Director to undertake specific projects. In 1969-1970, shatter the standing dead plant parts and are therefore an about 35 scientists were working on the Pawnee Site, effective agent of transfer from standing dead to litter, about 35 on the comprehensive network sites (some and (3) that root material is concentrated in the top 10 work on more than one site), and 8 in the central pro- cm of soil, differs from one grazing treatment to another gram. Most of the scientists receive support from the only slightly for the first 8 months of the calendar year, grassland program for only a few months each year. and decreases simultaneously in August under all About 70 graduate students are working in the program. treatments. There are a great number of species in the consumer trophic level. In our studies we have developed methods of determining intake, output, numbers, and biomass for RESEARCH METHODS AND PROGRESS all major species. For cattle we have already developed fistulation and fecal-collection methods. We have domes- ticated a small number of bison and antelope and we per- Research progress is reported for the Pawnee Site, form experiments with them. Of much significance in comprehensive network sites, modeling studies, and inter- studies of large herbivores has been the refinement of the disciplinary research coordination. water-intake method of estimating daily forage intake. Knowing body size, air temperature, and water intake, we can estimate daily forage intake to within 500 g per PAWNEE SITE animal per day. When small mammals and birds are studied, determin- Research progress at the intensive site is most logically ing forage intake and output by nondestructive methods covered in the direction of energy flow: abiotic, pro- is morc difficult. (Even estimating numbers and biomass ducer, consumer, and decomposer. Details are given in is more difficult.) Flushing transects have been used for Technical Reports 2, 3, 5-7, 9-23, 26-31, 33, 49, and estimating jackrabbits. Ground squirrels, kangaroo rats, 50-55. deer mice, and grasshopper mice have been trapped in The results of the abiotic studies have been used to grid systems. Age structure, breeding periods, andpreg-

12 nancy rates have been determined. Birdshave been inven- to be a complex, interrelated array of animal,plant, and toried by count and nest flush techniques. soil characteristics. Study of the ecosystem at only a few Dietary studies of small-mammal gut and bird crops points or considering only averages at many points would have been made by microscopic analysis of the contents. obscure many of these interrelationships. Our studies at the consumer level have been designed to The most obvious herbivorous insects on the Pawnee identify the feeding pathways. Site are the orthoptera. The orthoptera are consumed by The distribution of small mammals across the Pawnee the 13lined ground squirrel and the grasshopper mouse, Site is related to soils, intensity of grazing, vegetation by various small passerine birds, and by carnivorous in- patterns, and other factors. Thus far, however, norela- sects (e. g., robber flies). The orthoptera are animpor- tionship has been noted between intensity of grazing and tnat link in the transfer of energy andnutrients from the the number of grasshopper mice and 13lined ground producer compartment to these omnivorous feeders. squirrels that have been trapped. Because of the lower Over 30 species of orthopteran insects have been found density of kangaroo rats and deer mice, it is difficult to on the Pawnee Site. The most commonspecies are the determine whether there is a relationship between this grasshoppers. The dominant orthoptera on the Pawnee density and the intensity of grazing; apparently there is Site are not known tu be agricultural pests. Since invps- none. tigations began under this program, grasshoppers have There appear to be some relationships between soils invaded nearby cultivated lands, and the infestation was and rodents. Ord's kangaroo rats, northern grasshopper judged severe enough to warrant wide-scale spraying. The mice, and 13lined ground squirrels tend to avoid poorly pest species was not found on the Pawnee Site, whichin- drained, undifferentiated bottomland soils. Furthermore, dicates that certain natural dietary or other controls of kangaroo.rats prefer Renohill-Shingle soils to sandy pest species are present in native grasslands but not in loarns. Prairie deer mice appear to have no soil preference. cultivated lands. Both kangaroo rats and deer mice are much more Decomposers play the critical role of transferring or- abundant in specific habitats adjacent to the intensive- ganically bound nutrients to their available inorganic study pastures. Prairie deer mice are the most abundant states. A very sizable fraction, perhaps more thanhalf, of rodents in areas with especially tall grasses or thick shrub the ecosystem producer biomass, is not consumed by her- cover. Ord's kangaroo rat is probably the primaryrodent bivores but is directly decomposed by fungi and bacteria. biomass in sandy areas and in areas with loose soils and Fungi and bacteria account for most of the decom- low plant coverage. These areas are found along streams position, and attempts were made to determine the bio- on the edge of wheat or barley fields and inlow barrow mass of these organisms. Initial bacteria counts were pits near roads. made with a Petroff-Hauser counting chamber; in later Hares, perhaps because they are larger and more mo- studies a modification of the smear technique was used. bile, show predictable daily and seasonal preferences for Fungal biomass values were derived from estimates of assemblages of plants. Although the dispersion patterns mycelium length. These studies indicated that the fungal are species specific, there areconsiderable areas of over- biomass of soil was about 3 times that of bacterial bio- lap, particularly during the winter. Distribution patterns mass. For both bacteria and fungi,there was a decrease were investigated and revealed a very strongrelationship in biomass with soil depth; it was much more apparent with vegetational patterns. The black-tailed jackrabbit with fungi than with bacteria. A significant seasonal range occurred where there were standsof fourwing salt- trend also was observed; the biomass of both groups of bush, or in and near cultivated areas, or along the alluvial organisms decreased from July to September. plains of the Nunn series soil where western wheatgrass is The determination of microbial numbers or biomass the predominant grass species. White-tailed jackrabbits, involves the use of procedures that are laborious and on the other hand, were more common onupland soils time-consuming, and results are often difficult to inter- where shortgrasses (e. g., blue grama grass) are dominant. pret. To overcome some of these problems, we utilized Other examples of relationship between animal distri- adenosine triphosphate (ATP) measurements to give us butions and vegetation distributions were found for bird- microbial biomass. This method uses the luciferase en- nesting sites. Lark buntings apparently are more likely to zyme system. In the presence of ATP the systememits nest near taller vegetation, such as fourwingsaltbush and light, the amount being proportional to the available red three-awn. Leafhoppers are more common on the ATP. The relationship between ATP and bacterial bio- Nunn soil series occupied by western wheatgrass, and mass is extremely sensitive and linear over awide range grasshoppers are more common on soils occupied by blue of values, and it appears that the ATP procedure is satis- grama and buffalo grass. The lark bunting wasfound in factory for estimating biomass of soil bacteria. The very low numbers on pastures heavily grazedin summer, major shortcoming of the technique in its present form but large numbers of homed lark and McCown's longspur lies in the uncertainty of its ability to measure accurately were usually found on these pastures. Thus,there appears the fungal component of soil microflora.

11 13 We carried out other research to characterize the rate more deficient in energy than the grass-covered areas. of iiecomposition by fungi and bacteria, using filter The results indicate that fixation by free-living bacteria paper buried at different times during the fall of 1969. probably is not an important source of nitrogen on the These studies indicated, as ex9ected, that plant litter and grassland. There may, however, be localized areas with cellulose disappeared most rapidly during times of warm,. high levels of energy supplying material, coupled with wet weather; during periods of very cool weather there anerobic conditions, that could result in substantial rates was almost no loss of these materials. The techniques of fixation. On the Pawnee Site, these areas are confined will be refined and expanded so that a better quantitative to the intermittent lakes. picture of the decomposition rate of below-ground plant material and plant litter can be obtained. Preliminary work on phyllosphere and litter bacteria COMPREHENSIVE NETWORK SITES involved mostly the selection of suitable nutrient agar for optimum development of these microbial popula- Three meetings were held to organize the comprehen- tions. The results indicated that a large proportion of the sive network: bacteria associated with living (green) plant tissue re- At the first meeting, held in July 1969, the sites quired organic growth factors and that a small proportion were selected-10 sites in 9 western statesal:d a general of the bacteria associated with dead plant tissue required plan of trophic level investigation was worked out for them. These findings suggest that the living plant provides each. The meeting was attended by representatives of growth factors required by phyllosphere microorganisms each site, subject-area specialists, and some of the scien- and show that direct decomposition of plant material in- tific coordinators from the central grassland biome office. volves a wide range of bacteria. The second meeting was held in November 1969 in Soil respiration is an index of the physiological activ- conjunction with the annual meeting at the Pawnee Site. ities of the soil organism and of the turnover rate of the During the first 2 days, investigators from the Pawnee organic matter in the soil. Rates of CO2 evolution from Site gave reports on progress achieved in aseason of field soil cores w^re measured in the laboratory. Soil cores work. These reports enabled the investigators from the were divided into sections taken from depths of 0-3, 3-6, network sites to anticipate problems and to adjust their 6-10, 10-20, 20-50, 50-70, and 70-90 cm. During the research designs. In the remaining 11h days, discussion first day of incubation, CO2 evolution was about the was devoted to research at the network sites. After a re- same for the various upper-core sections. There was a view of important variables in each trophic levei, the high rate of CO2 evolution in the sections from the depth group divided into workshops on herbage dynamics, ver- of 50-70 cm, indicating that the aeration condition in the tebrates, invertebrates, and microbiology. In the work- extracted cores is considerably different from that of the shops, plans were made regarding variables to be evalu- soil in situ. Future investigations of CO2 evolution will ated, desired precision, and problems requiring further utilize more frequent sampling in the field, and there will consideration. be concurrent measurements of soil temperature and soil A small group representing the various specialties moisture at various depths. Also, laboiatory investigations met in December 1969. They reviewed plans for studying will utilize similar techniques where soil moisture and soil trophic levels and discussed changes in plans that had temperature are held at constant values. been made since the November meeting. Nitrogen must be replaced constantly in the ecosystem The plans and decisions developed during these meet- by various fixation mechanisms, and the fixation rateson ings were summarized in a manual of field procedures for the predominantly grass-covered sites are very, very low. the comprehensive network (Technical Report 35). The The acetylene reduction technique, because of its high variables and measurements required were organizedon sensitivity, is used (Technical Report 7). Cores from an field data sheets, which were printed and distributed for area vegetated with buffalo grass (Buchlod dactyloides) the use of all field investigators. The forms provide for and soil cores from the bare-soil area of an intermittent consistent coding of all data and for organizing them for lake were compared while they were saturated with mois- keypunching. ture. Cores from buffalo grass areas fixed 5 g per hectare The 10 network sites selected for investigation in the per day, and cores from bare-soil areas fixed 1.2 g per United States fit into a design for comparing grassland hectare per day. habitats in various parts of North America (Figure 3). Addition of a source of soluble energy (sucrose)re- Other sites being considered are the Matador Site in sulted in a greater increase in nitrogen fixation rateson Saakatchewan, Canada, and the Campana Site in Chihua- the (bare-soil) cores than on the sodded-grass cores. This hua, Mexico. Investigation of the grasslands biome in perhaps can be interpreted to mean that the bacteriaor three North American nations provides unequalledop- algal populations in the intermittent lake situationsare portunities for comparative analyses. Such comparisons more suited for nitrogen fixation. But these areas are are essential to understanding the interrelationships of

12 14 ecosystem compartments in the function of the total tional methoe.s for evaluation of effective area. ecosystem. A comparative evaluation of three methods of sam- A certain minimum set of data is required from a site pling insect populations was made before field sampling in order for it to contribute to the objectives of the com- began. Sampling by sweep net grossly underestimated prehensive network project. Characterization of the driv- numbers of insects. Sampling by a vacuum net overesti- ing forces in the abiotic compartment of the ecosystem mated groups that jump or fly weakly when flushed by requires measurement of total and net radiation, precip- the apparatus and underestimated other groups. The itation, wind, atmospheric moisture, the temperature quick trap, which encloses an area that is then thoroughly gradient above and below the surface of the ground, the collected by a vacuum device, gave the most accurate gradient of soil moisture, and soil heat flux (see Technical estimates. Report 35). Quantitative characterization of primary pro- The length of the growing season varies considerably duction requires measurement of above-ground and among sites, and the initiation of field work varies ac- below-ground plant biomass and litter. Consumer popula- cordingly. Eight sampling periods have been completed tions, both vertebrate and invertebrate, must be quanti- for vegetation studies at the Osage Site in Oklahoma. fied at various times during the season. Decomposer ac- During the same length of time thcre have been four sam- tivity must be quantified. pling periods for below-ground biomass, which is deter- Ten battery-powered micrometeorological packages mined from soil cores. Two estimates of small mammal have been designed and built to record the abiotic vari- populations, one before and one after the breeding sea- ables mentioned here. The package includes a recording son, have been made for five of the sites, and one esti- unit that will read hourly from 13 sensors and record re- mate has been made for two additional sites. Evaluation sults on magnetic tape. The tape will be returned monthly of the avian populations has been completed at eight of to the central grassland biome office, and the results will the sites. be summarized, returned to the investigators, and added to the information storage and retrieval system. Appro- priate summaries of the data will be distributed to the MODE LI NG RESU LTS investigators and scientific coordinators. A meeting of the investigators and graduate students Since initiation of the program in 1968, we have pon- sampling small mammals on the comprehensive network dered the question of how best to organize the modeling sites was held in April 1970 at the Jornada Site in New activity, for which manpower was limited. In the autumn Mexico. Scientists from three universities and one federal of 1969, we organized a group to develop preliminary agency attended. The investigators participated in a field- models. This group included three scientists in ecological sampling exercise to gain experience in solving logistic modeling and analysis, three persons working primarily problems that would be met in the network. This success- in an interactive mode between the analysts and the ex- ful effort stimulated similar meetings for other disciplin- perimentalists interested in getting their ideas and data ary groups. For example, a meeting of scientists sampling into models. Much of our initial activity was devoted to invertebrates was held at the Pantex Site in Texas. conceptual and methodological reviews of mathematical An experimental evaluation of the effectiveness of the modeling of ecological systems.* small-mammal sampling program being followed in the The systems analysis group has developed several spe- grasslands biome was conducted in July in southern cific subsystem or total-system simulation models: Nevada. A group of investigators from the University of An example of a subsystem model is a model of California at Los Angeles have regularly censused the ro- bird population and biomass dynamics, patterned after dent populations for several years , using 8-hectare en- the Pawnee Site situation (Technical Report 3). This closures. Personnel from the subprogram concerned with model has two differential equations that, when solved, the desert biome and from the Savannah River Ecology give the dynamics of bird numbers and biomass. Coeffi- Laboratory participated. IBP sampling methods were ap- cients in the model are based on data for the lark bunting. plied in the enclosures, and the resulting population esti- A more complex analytical model is concerned mates were compared with the true values. Certain with the population dynamics of small mammalian her- methods of computing population estimates (e.g., those bivores (Technical Report 4). assuming a geometric frequency distribution of recapture) A simplified total-ecosystem energy-flow model were excluded as a result of the studies. Other methods was developed in 1968 as a preliminary effort for a total gave more realistic values. The procedures followed in the field also permitted determination of the locations *See references to the following articles in Appendix A (p. 14): "Model Structure for a Grassland Ecosystem," by Bledsoe and from which animals came as they moved toward the traps: Jameson (1969), and "Implementing the Ecosystem Concept in Therefore, the effective area of the trapping grid could be Training in the Natural Resource Sciences," by Van Dyne (1969). evaluated empirically for comparison with the computa- see also Technical Report 32.

15 system. This model contained generalized plants, animals, ably needed in this kind of investigation. Perhaps 80 to and microbes with coefficients patterned specifically 90 percent of the participants should work on individual after warm-season shortgrasses as producers, antelopes as subprojects requiring about 25 percent of their time. The herbivores, and coyotes as predators. In this model an in- others should devote their efforts to the overall program cremental approach was used to update changes in bio- and, for about 75 percent of their time, should share mass of the animal and plant components in a system. work on the following tasks: (1) regional, national, and The driving forces utilized in this model were the clima- international development, which includes describing the tological records from the Pawnee Site. program to, and evaluating it with, scientific, government, An initial complex macromodel, composed of 46 and industrial groups; (2) scientific coordination, which differential equations, was developed early in 1970. This includes program management and administration, organ- highly nonlinear but deterministic model has four driv- izing and conducting meetings, developing reports and ing variables and 46 principal systems variables. proposals, and planning and evaluation; (3) scientific in- The systems analysis group also was involved ina va- tegration and synthesis, which includes working with riety of specialized mathematical analyses. For example, participants in Afferent trophic areas of specialization. the thy-weight rank method was examined. We inves- The program scientists would also pursue individual re- tigated its theory, distributional properties of parameters search in their areas of expertise. derived from the model, and limitations of the technique Thus, we need a core of scientists dedicated to the for determining botanical composition of plant commu- overall program. But our plan also encourages the con- nities. Another group of specialized models reviewed in- tributions and cooperation of a much larger number of cludes those for inferring evaporation from meteorolog- scientists from universities and federal laboratories. Their ical measurements (Technical Report 47). cooperation is necessary for a more complete understand- To provide techniques needed for future stochastic ing of the entire ecosystem. and statistical work, we developed special packages that included distribution generators for the F statistic and the Chi-square statistic. Also, because in stochastic simulations several parameters may need to be varied simul- APPENDIX A: PUBLICATIONS taneously, we developed a generator for multivariate normal distribution. We developed computer programs Technical Reports* for solving differential equations, for optimization, and 1 Jameson, D. A., and R. E. Bement. 1969. General descrip- for statistical analyses, then published technical reports tion of the Pawnee Site. 32 p. to enable participants to make wide use of them (Tech- 2Bartos, D., and J. Hughes. 1969. Preliminary methodology nical Reports 34 and 46). Analytical personnel havepar- and results for root biomass sampling on the Pawnee Site. ticipated in field-based workshops in which special atten- 20 p. 3Swartzman, G. S. 1969. A preliminary bird population tion was given to sampling problems related to estimating dynamics and biomass model. 16 p. populations of mammals and insects. 4 Gross, J. E., and C. J. Walters. 1969. Summary report on initial small-herbivorous-mammal modeling efforts. 57 p. 5Striffler, W. D., and F. M. Smith. 1969. Pawnee Site microwatersheds: selection description and instrumentation. 29 p. INTERDISCIPLINARY RESEARCH 6Galbraith, A. F. 1969. Soil water study of a shortgrass prairie ecosystem, Pawnee Site. 51 p. 7Reuss, J. 0., and P. W. Copley. 1969. Soil nitrogen investi- We are making some important changes in the struc- gations, Pawnee Site. 13 p. ture of the grassland program. Some unique features are 8Jameson, D. A., and L. G. Nell. 1970. Memoranda of agree- becoming clear. ment and procedures for working on federal lands of the First, a great deal of time must be spent in program USDA. 53 p. 9Fisser, H. G. 1969. Preliminary report of methodology and development, organization, and scientific public relations. results for analysis of plant pattern subproject research on For example, since 1968, the Grasslands Biome Director the Pawnee Site. 65 p. has given more than 20 lectures in which he has described, 10 Hyder, D. N., K. L. Knox, and R. E. Bement. 1969. Meta- for legislative, scientific, and industrial groups, the con- bolic components of cattle: water-soluble tracers for determining water turnover and partitioning by cattle. 32 p. cepts and operation of the program. 11Rice, R. W., D. R. Cundy, and P. R. Weyerts. 1969. A com- Second, to maintain the necessary coordination, ex- parison of the esophageal fist--ta with rumen samples for change of information and data, and interaction among the determination of the botanical and chemical composi- the participants, it appears that 0.5 to 1.0 man-month is tion of the diet of herbivores. 13 p. required for each senior participant simply to attend various types of meetings, seminars, discussion group ses- *The number preceding each item is the report number. Copies sions, and workshops each year. of these reports may be obtained by writing to the author or to the Grasslands Biome Director (George M. Van Dyne). Supplies Third, two major types of senior participants are prob- are limited.

16 Herbel, C. H., and R. D. Pieper. 1970. Comprehensive net- 12Rice, R. W., and M. Vavra. 1969. Botanical species of plants 43 eaten and intake of steers grazing light, mc Jium, and heavy work site description, Jornada. 21 p. use shortgrass range. 18 p. 44Risser, P. G. 1970. Comprehensive network site description, Osage. 5 p. 13Nagy, J. G., K. L. Knox, and D. E. Wesley. 1969. Progress de- report 1BP antelope project, Pawnee Site. 18 p. 45 Huddleston, E. W. 1970. Comprehensive network site scription, Pantex. 12 p. 14Hansen, R. M., J. T. Flinders, and B. R. Cavender. 1969. Dietary and energy relationships of jackrabbits at the 46Bledsoe, L. J. 1970. ODE: numerical analysis for ordinary differential equations. 42 p. Pawnee Site. 43 p. 47 Nunn, J., L. J. Bledsoe, and R. D. Burman. 1970. Models 15Flake, L. D. 1969. A study of rodents in northeastern for inferring evaporation from meteorological measurements. Colorado. 29 p. Gross, J. E. 1969. Jackrabbit demographic and life history 20 p. 16 48Huddleston, E. W., C. R. Ward, R. E. Howard, and L. G. studies, Pawnee Site. 8 p. Richardson. 1969. Some contributions to the study of grass- 17Bertolin, C and J. Rasmussen. 1969. Preliminary report on Grass- lands insect populations. 9 p. the study of tho precipitation on the Pawnee National 49 Herrmann, S. J., J. W. Lavelle, and J. A. Seilheimer. 1970. land. 34 p. Aquatic primary productivity and physical-chemical lim- 18Cavender, B. R., and R. M. Hansen. 1970. The microscope nology on the Pawnee Site. 19 p. method used for herbivore diet estimates and botanical 1970. analysis of litter and mulch at the Pawnee Site. 9 p. 50 Thatcher, T. 0., Grace Inyamah, and J. E. Mitchell. Sampling insect populations by sweep net on the Pawnee 19Van Horn, D. H. 1969. Dry weight biomass data for four abundant grasshopper species of the Pawnee Site. 6 p. Site, 10 p. 51Hansen, R. M., and Barbara Cavender. 1970. Assimilation 20Lavigne, R. J., -tind L. E. Rogers. 1970. Effect of insect predators and parasites on grass feeding insects, Pawnee Site. rates of small mammal herbivores. 7 p. 38 p. 52Clark, F. E. 1970. The microbial component of the ecosys- 21Doxtader, K. G. 1969. Microbial biomass measurements at tem. 14 p. the Pawnee Site: preliminary methodology and results. 16 p. 53 Cwik, M. J. 1970. Identification of insects and density de- 22 Mayeux, J. V., and E. A. Jones. 1969. Bacterial ecology of terminations of the stomach contents of small mammals. grassland soils, Pawnee Site. 13 p. 10 p. 54 Wolff, D. N. 1970. Grassland infiltration phenomena.125 p. 23Christensen, M., and A. M. Scarborough. 1969. Soil microfungi investigations, Pawnee Site. 18 p. 55Streeter, C. L. 1970. Standardized processing and storage 24 Wright, R. G. [compiler] . 1970. Scientific personnel partic- scheme for samples collected for IBP Grassland Ecology Re- ipating in the Grassland Biome study, June 1968 through search Laboratory. 5 p. January 1970.278 p. 56Hendricks, Barbara J. 1970. Style and format of technical report. 42 p. 25Robinson, R. D. 1970. IBP Grasslands Biome budget program. 10 p. 26Ryder, R. A. 1969. Diurnal raptors on the Pawnee Site. 16 p. Journal Articles, Books, Parts of Books, and Theses 27Marti, C. D. 1969. Some comparisons of feeding ecology of four species of owls in north-central Colorado. 21 p. Alexander, M. 1969. Soil decomposers, p. 403-409. In R.L. Dix 28Giezentanner, J. B., and R. A. Ryder. 1969. Avian distribu- and R. G. Beidleman [ed.] The grassland ecosystem: a pre- tion and population fluctuations, Pawnee Site. 29 p. liminary synthesis. Range Science Department, Science 19Baldwin, P. H., J. D. Butterfield, and P. D. Creighton. 1969. Series No. 2. Colorado State Univ., Fort Collins. Summer ecology of the lark bunting, Pawnee Site. 37 p. Bement, R. E. 1969. Dynamics of standing deadvegetation on R. G. 30Franklin, W. T. 1969. Mineralogy of representative soils at the shortgrass plains, p. 221-224. /n R. L. Dix and Beidleman [ed.] The grassland ecosystem: a preliminary the Pawnee Site. 10 p. 2. Moir, W. H. 1969. Photosynthesis of shortgrasses under synthesis. Range Science Department, Science Series No. 31 Colorado State Univ., Fort Collins. field conditions. 31 p. digital Swartzman, G. [coordinator] . 1970. Some concepts of Bledsoe, L. J., and G. M. Van Dyne. 1969. Evaluation of a 32 computer method for analysis of compartmentalmodels of modeling. 142 p. ecological systems. Oak Ridge Nat. Lab. TM-2414.60 p. 33 Uresk, D., and P. L. Sims. 1969. Preliminary methodology Bledsoe, L. J., and D. A. Jameson. 1969. Model structurefor a and results for aboveground herbage biomass sampling on grassland ecosystem, p. 410-437. in R. L. Dix and R. G. the Pawnee Site. 13 p. Beidleman [ed.] The grassland ecosystem: a preliminary 34Swift, D. M. [coordinator] . 1970. Current generalized synthesis. Rang( Science Department, Science Series No.2. computer programs used in Grassland Biome analyses. Colorado State Univ., Fort Collins. 286 p. Blocker, H. D. 1969. The impact of insects as herbivoresin 35French, N. R. 1970. Field data collection procedures for grassland ecosystems, p. 290-299. In R. L. Dix andR. G. the comprehensive network 1970 season. 35 p. Beidleman [ed.] The grassland ecosystem: a preliminary 36Rickard, W. H., and T. P. O'Farrell. 1970. Comprehensive synthesis. Range Science Department, Science Series No.2. network site description, arid lands ecology. 5 p. Colorado State Univ., Fort Collins. 37Morris, M. S. 1970. Comprehensive network site description, Bt,rzlaff, D. F. 1969. The role of the abiotic factors in the Bison. 23 p. structure and function of the grassland ecosystem, p. I17- 38Collins, D. 1970. Comprehensive network site description, 123. /n R. L. Dix and R. G. Beidleman [ed.] Thegrassland Bridger. 10 p. ecosystem: a preliminary synthesis. Range ScienceDepart- 39Lewis, J. K. 1970. Comprehensive network site description, ment, Science Series N. 2. ' !olorado State Univ.,Fort Cottonwood. 26 p. Collins. 40 Whitman, W. C. 1970. Comprehensive network site descrip- Butterfield, J. D. 1969. Nest site requirements of the lark tion, Dickinson. 15 p. buting in Colorado. M.S. thesis. Colorado State Univ., Fort 41Tomanek, G. W. 1970. Comprehensive network site descrip- Collins. 59 p. tion, Hays. 6 p. Clark, F. E. 1969. The microflora of grassland soils and some 42 Heady, H. F. 1970. Comprehensive network site description, microbial influences on ecosystem functions, p. 361-376. Hopland. 11 p. In R. L. Dix and R. G. Beidleman [ed.] The grassland eco-

15 system: a preliminary.synthesis. Range Science Department, energy flux, water flux, and nutrient flux in grasslan6 eco- Science Series No. 2. Colorado State Univ., Fort Collins. systems, p. 197-220.Th R. L. Dix and R. G. Beidieman [ed.] Collins, D. D. 1969. Macroclimate and the grassland ecosystem, The grassland ecosystem: a preliminary synthesis. Range p. 29-39. /n R. L. Dix and R. G. Beidlemaa [ed.] The grass- Science Department, Science Series No. 2. Colorado State land ecosystem: a preliminary synthesis. Range Science De- Univ., Fort Collins. partment, Science Series No. 2. Colorado State Univ., Fort Lavelle, J. W., J. A. Seilheimer, N. L. Osborn, and S. J. Herrmann. Collins. 1969. A preliminary study of three lentic communities on Dix, R. L. 1969. A history of the North American grasslands as the Pawnee National Grasslands, p. 308-315. In R. L. Dix and related to ecosystem studies 27. In R. L. Dix and R. G. R. G. Beidleman [ed.] The grassland ecosystem: a prelimi- Beidleman [ed.] The grastand ecosystem: a preliminary nary synthesis. Range Science Department, Science Series synthesis. R ange Science Department, Science Series No. 2. No. 2. Colorado State Univ., Fort Collins. Colorado State Univ., Fort Collins. Lewis, J. K. 1970. Primary producers in grassland ecosystems, Doxtader, K. G. 1969. Estimation of microbial biomass in soil p. 241-287. In R. L. Dix and R. G. Biedleman [ed.] The on the basis of adenosine triphosphate. Bacteriological pro- grassland ecosystem: a preliminary synthesis. Range Science ceedings abstracts of 69th annual meeting, American Society Department, Science Series No. 2 (supplement). Colorado for Microbiology. State Univ., Fort Collins. Everson, A. D. 1969. Replacement of native plant communities Marti, C. D. 1969. Renesting by barn owls and great horned with introduced communities and its impact on ecosystem owls. Wilson Bull. 81:467-468. function, p. 261-267. /n R. L. Dix and R. G. Beidleman Marti, C. D. 1969. Some comparisons of the feeding ecology of [ed.] The grassland ecosystem: a preliminary synthesis. four owls in north-central Colorado. Southwest. Natur. Range Science Department, Science Series No. 2. Colorado 14:163-170. State Univ., Fort Collins. Moir, W. H. 1968. Prairie rebirth. Science 162:1312. Fisser, H. G. 1969. Plant pattern and distribution in ecosystems Moir, W. H. 1969. Energy fixation and the role of primary pro- and relationships to function, p. 183-196. /n R. L. Dix and ducers in energy flux of grassland ecosystems, p. 125-147. Ft . G. Beidleman [ed.] The grassland ecosystem: a prelimi- In R. L. Dix and R. G. Beidleman [ed.] The grassland eco- nary synthesis. Range Science Department, Science Series system: a preliminary synthesis. Range Science Department, No. 2. Colorado State Univ., Fort Collins. Science Series No. 2. Colorado State Univ., Fort Collins. Glover, F. A. 1969. Birds in grassland ecosystems, p. 279-289. Nagy, J. G., T. A. Barber, and A. E. McChesney. 1969. Closter- In R. L. Dix and R. G. Beidleman [ed.] The grassland eco- dium perfringens enterotoxemia in hand-reared antelope. J. system: a preliminary synthesis. Range Science Department, Wildlife Management 33:1032-1033. Science Series No. 2. Colorado State Univ., Fort Collins. Paris, 0. H. 1969. The function of soil fauna in grassland eco- Gross, J. E. 1969. The role of small herbivorous mammals in systems, p. 331-360. /n R. L. Dix and R. G. Beidleman [ed.] the functioning of the grassland ecosystem, p. 268-278. In The grassland ecosystem: a preliminary synthesis. Range R. L. Dix and R. G. Beidleman [ed.] The grassland ecosystem: Science Department, Science Series No. 2. Colorado State a preliminary synthesis. Range Science Department, Science Univ., Fort Collins. Series No. 2. Colorado State Univ., Fort Collins. Pieper, R. D. 1969. The role of consumers in a grassland eco- Hansen, R. M., and .1. T. Flinders. 1969. Food habits of North system, p. 316-329. In R. L. Dix and R. G. Beidleman [ed.] American hares. Range Science Department, Science Series The grassland ecosystem: a preliminary synthesis. Range No. 1. Colorado State Univ., Fort Collins. 18 p. Science Department, Science Series No. 2. Colorado State Haug, P. T. 1969. The International Biological Program-the Univ., Fort Collins. grassland ecosystem analysis. J. Colorado-Wyoming Academy Pochop, L. 0.1969. Dynamics of the atmosphere in the grass- of Science 6:4-5. Abstract, 45th annual meeting, Southwest- land ecosystem, p. 89-100. In R. L. Dix and R. G. Beidleman ern and Rocky Mountain Division of the American Associa- [ed.] The grassland ecosystem: a preliminary synthesis. Range tion for the Advancement of Science, and the 40th annual Science Department, Science Series No. 2. Colorado State meeting, Colorado-Wyoming Academy of Science. Univ., Fort Collins. Haug, P. T. 1970. Succession on old fields: a review. M.S. thesis. Porter, L. K. 1969. Nitrogen in grassland ecosystems, p. 377- Colorado State Univ., Fort Collins. 473 p. 402. /n R. L. Dix and R. G. Beidleman [ed.] The grassland Haug, P. T., G. M. Van Dyne, and R. M. Hansen [ed.]. 1969. ecosystem: a preliminary synthesis. Range Science Depart- Dietary competition among herbivores-papers from a joint ment, Science Series No. 2. Colorado State Univ., Fort graduate seminar between wildlife biology and range science. Collins. Colorado State Univ., Fort Collins. 121 p. Post, G. 1969. The role of diseases and parasites in a grassland Hughes, J. H. 1969. An evaluation of the dry weight rank ecosystem, p. 300-306. In R. L. Dix and R. G. Beidleman method of determining species composition of plant com- [ed.] The grassland ecosystem: a preliminary synthesis. Range munities. M.S. thesis. Colorado State Univ., Fort Collins. Science Department, Science Series No. 2. Colorado State 111 p. Univ., Fort Collins. Hyder, D. N. 1969. The impact of domestic animals on the Risser, P. G. 1969. Competitive relationships among herbaceous functien and structure of grassland ecosystems, p. 243-260. grassland plants. Bot. Rev. 35:251-284. In R. L. Dix and R. G. Beidleman [ed.] The grassland eco- Scarborough, A. M. 1970. The soil microfungi of a Colorado system: a preliminary synthesis. Range Science Department, grassland. M.S. thesis. Univ. Wyoming. 68 p. Science Series No. 2. Colorado State Univ., Fort Collins. Shubert, M. L. 1969. The nature and importance of competition Inyamah, G. C. 1969. Population trends of insects on Melilotus between woody and herbaceous plants in a grassland eco- officinalis (L) Lam. in the Pawnee Grassland. M.S. thesis. system, p. 172-182. In R. L. Dix and R. G. Beidleman [ed.] Colorado State Univ., Fort Collins. 69 p. The grassland ecosystem; a preliminary synthesis. Range Kelly, M. J., P. A. Opstrup, J. S. Olson, S. I. Auerbach, and Science Department, Science Series No. 2. Colorado State G. M. Van Dyne. 1969. Models of seasonal primary produc- Univ., Fort Collins. tivity in eastern Tennessee Festuca and Andropogon eco- Striffler, W. D. 1969. The grassland hydrologic cycle. p. 101- systems. Oak Ridge National Laboratory TM-4310.296 p. 116. /n R. L. Dix and R. G. Beidleman [ed.] The grassland Kline, J. R. 1969. Soil chemistry as a factor in the function of ecosystem: a preliminary synthesis. Range Science Depart- grassland ecosystems, p. 71-88. In R. L. Dix and R. G. ment, Science Series No. 2. Colorado State Univ., Fort Beidleman [ed.] The grassland ecosystem: a preliminary Collins. synthesis. Range Science Department, Science Series No. 2. Terwilliger, C., Jr. 1969. Physical properties of grassland soils Colorado State Univ., Fort Collins. and their influence on primary productivity, p. 65-70. /n Knight, D. H. 1969. Some influences of vegetation structure on R. L. Dix and R. G. Beidleman [ed.] The grassland ecosys-

16 18 tern: a prelhninary synthesis. Range Science Department, mer growth in Bouteloua gracilis. American Society of Range Science Series No. 2. Colorado State Univ., Fort Collins. Management Annual Meeting, Denver, Colo. Feb. 9-12. Thomas, B. 0., R. E. Cameron, and J. D. Holmes. 1970. The Christensen, M. ! 969. Major fungal populations on the Pawnee importance and role of arnphibians and reptiles in grassland Site. U.S. Grassland Biome Seminar, Univ. Wyoming. ecosystems, p. 307-323. In R. L. Dix and R. G. Beidleman Cook, C. W. 1970. Thc ecosystem concept in an undergraduate [ed.] The grassland ecosystem: a preliminary synthesis. zurriculum in range science. American Society of Range Range Science Department, Science Series No. 2 (supple- Management Annual Meeting, Denver, Colorado. Feb. 9-12. ment). Colorado State Univ., Fort Collins. Cook, C. W. 1970. Ecosystem approach to teaching range sci- Tomanek, G. W. 1969. Dynamics of mulch layer in grassland ence. American Society of Range Management Annual Meet- ecosystems, p. 225-240. In R. L. Dix and R. G. Beidleman ing, Denver, Colo. Feb. 9-12. [ed.] The grassland ecosystem: a preliminary synthesis. Dick-Peddie, W. A., and W. H. Moir. 1970. Vegetation of the Range Science Department, Science Series No. 2. Colorado Organ Mountains, N. Mex. Range Science Department, Sci- State Univ., Fort Collins. ence Series No. 4. Colorado State Univ., Fort Collins. 28 p. Van Dyne, G. M., F. N. Glass, and P. A. Ostrup. 1968. Develop. Edwards, P., and D. Striffler. 1970. The central data acquisition men t and use of capacitance meters to measure standing crop system on the Pawnee Site. U.S. Grassland Biome Seminar, of herbaceous vegetation. Oak Ridge Nat. Lab. TM-2247. Univ. Colorado. 47 p. Fraley, L. 1969. Studies with the 8700 Curie radiation source Van Dyne, G. M. 1969. MeasuUng quantity and quality of the on the Pawnec Site. U.S. Grassland Biome Seminar, Southern diet of large herbivores, p. 54-94. In F. B. Golley and H. K. Colorado State Coll. Duechner [ed.] A practical guide to the study of the pro- Free, J. C., R. M. Hansen, and P. L. Sims. 1970. Methods for ductivity of large herbivores. Blackwell Scientific Publications, estimating dry-weight composition in diets of large herbi- Oxford. vores. American Society of Range Management Annual Van Dyne, G. M. [ed.] . 1969. The ecosystem concept in natural Meeting, Denver, Colo. Feb. 9-12. resource management. Academic Press, Inc., New York. 383 p. Hansen, R. M., and J. C. Free. 1970. tviinimal observations Van Dyne, G. M. 1969. A plea for fewer but more-significant needed to determine dry-weight compo'' ion in herbivore digits. J. Range Manage. 22:52-53. diets by microscopic methods. American Society of Range Van Dyne, G. M. 1969. The problem, need, and mode of sny- Management Annual Meeting, Denver, Colo. Feb. 9-12. thesis in the grassland biome study, p. 1-2. In R. L. Dix and Haug, P. T. 1970. Analysis of the American grassland biome: R. G. Beidleman [ed.] The grassland ecosystem: a preliminary An overview of an integrated research program. American synthesis. Range Science Department, Science Series No. 2. Society of Range Management Annual Meeting, Denver, Colorado State Univ., Fort Collins. Colo. Feb. 9-12. Van Dyne, G. M. 1969. Some mathematical models of grasslands Herrmann, S., J. Lavelle, and J. Seilheimer. 1969. Aquatic ecosystems, p. 3-26. In R. L. Dix and R. G. Beidleman [ed.] studies on the Pawnee Site. U.S. Grassland Biome Seminar, The grassland ecosystem: a preliminary synthesis. Range Colorado State Coll. Science Department, Science Series No. 2. Colorado State Hyder, D. N. 1970. The leaf replacement potential of grasses. Univ., Fort Collins. American Society of Range Management Annual Meeting, Van Dyne, G. M. 1969. Implemendng the ecosystem concept in Denver, Colo. Feb. 9-12. training in the natural resource sciences, p. 327-367. In Jameson, D. A. 1970. A mathematical model of a grassland eco- G. M. Van Dyne [ed.] The ecosystem concept in natural re- system. American Society of Range Management Annual source management. Academic Press, Inc., New York 383 p. Meeting, Denver, Colo. Feb. 9-12. Van Dyne, G. M. 1969. Grasslands management, research, and Jameson, D. A. 1970. Modelling and systems analysis in range training vieweei in a systems context. Range Science Depart- ;cience. American Society of Range Management Annual ment, Science Series No. 2. Colorado State Univ., Fort Meeting, Denver, Colo. Feb. 9-12. Collins. 35 p. Jameson, D. A. 1969. Land management policy and the develop- Van DI ne, G. M. 1970. Animals of grassland ecosystems. En- ment of ecological concepts. USDA Forest Service Seminar, cyclopedia of science and technology. McGraw-Hill, Inc., Great Falls, Mont. Feb. 9. New York. Jameson, D. A. 1969. The biological function of land. Montana Van Dyne, G. M., W. E. Frayer, and L. J. Bledsoe. 1970. Optimi- Conservation Council Meeting, Billings, Mont. April 19. zation techniques and problems in the natural resource sci- Marti, C. D. and R. A. Ryder. 1969. Food habits of four grass- ences, p. 95-124. In Studies in optimization. Society for land inhabiting owls. Joint annual meeting of the South- Industrial and Applied Mathematics, Philadelphia. western and Rocky Mountain Division of the American Ward, R. T. 1969. The nature and significance of eco-genetic Association for the Advancement of Science and the variations in ecosystems, p. 148-152. In R. L. Dix and R. G. Colorado-Wyoming Academy of Science. Colorado Sprhigs, Beidleman [ed.] The grassland ecosystem: a preliminary syn- Colo. May 7-10. thesis. Range Science Department, Science Series No. 2. Miller, L. D. 1970. Automatic interpretation of remotely pro- Colorado State Univ., Fort Collins. cured data for vegetation inventories. American Society of Whitman, W. C. 1969. Microclimate and its importance in grass- Range Management Annual Meeting, Deaver, Colo. Feb. land ecosystems, p. 40-64. In R. L. Dix and R. G. Beidleman 9-12. [ed.] The grassland ecosystem: a preliminary synthesis. Nunn, J., and R. Weeks. 1970. The movable meteorological data Range Science Department, Science Series No. 2. Colorado acquisition system at the Pawnee Site. U.S. Grassland Biome State Univ., Fort Collins. Seminar, Colorado State Coll. Ryder, R. A. 1969. Birds, bison, and beef. 68th annual meeting of the Nebraska Ornithologists Union, North Platte Station, Neb. May 16-18. Ryder, R. A. 1970. IBP grassland biome, studies. Colorado APPENDIX B: PRESENTATIONS Junior Academy of Science Meetim, Cherry Creek, Colo. Feb. 2. Bement, R. E. 1970. Leaf-weight management on blue grama Ryclor, R. A. 1970. Censuses of grassland birds in Colorado range. American Society of Range Management Annual Meet- under the IBP. Third International Bird Census Conference, ing, Denver, Colo. Feb. 9-12. Oosterbeek, The Netherlands. Aug. 24-28. Bledsoe, L. J. 1969. Modeling of ecological systems. Rocky Ryder, R. A., and D. A. Cobb. 1969. Birds of the Pawnee Mountain Simulation Council Meeting, Denver, Colo. May 28. National Grassland in northern Colorado. Joint annual Burzlaff, D. F., and J. L. Stubbendieck. 1970. Nature of phyto- meeting of the Southwestern and Rocky Mountain Division

17 of the American Association to: the Advancement of Science State University, Fort Collins, Colo. 80521 (biometrical and the Colorado-Wyoming Academy of Science, Colorado services; modeling) Springs, Colo. May 7-10. W. T. Franklin, Agronomy Department, Colorado State Univer- Ryder, R. A ., J. B. Giezentanner, and C. Marti. 1970. Birds of sity, Fort Collins, Colo. 805 21 (soil mineralogy) the Pawnee Site: population dynamics and birds of prey. US. N. R. French, Natural Resource Ecology Laboratory, Colorado Grassland Biome Seminal, Colorado State Coll. State University, Fort Collins, Colo. 80521 (Comprehensive Sims, P. L., D. N. Hyder, L. J. Ayuko, and R. K. Lang. 1970. Network Director) Developmental morphology of four warm-season grasses. J. W. Gerdeman, Department of Plant Pathology, University of American Society of Range Management Annual Meeting, Illinois, Urbana, Ill. 61803 (soil microbiology) Denver, Colo. Feb. 9-12. R. W. Gorden, Department of Biology, Texas Technological Striffler, W. D. 1970. Hydrologic research in the 1BP grassland University, Lubbock, Tex. 79409 (soil microbiology) program. American Society of Range Management Annual J. E. Gross, Cooperative Wildlife Units, Colorado State Univer- Meeting, Denver, Colo. Feb. 9-12. sity, Fort Collins, Colo. 805 21 (small mammals) Tomanek, G. W. 1969. Dynamics of the mulch layer in the R. M. Hansen, Rarer Science Department, Colorado State Uni- grassland ecosystem. National meeting of the Southwestern versity, Fort Collins, Colo. 80521 (small mammals', Association of Naturalists, Tempe, Ariz. C. L. Hanson, Agricultural Research Service, U.S. Department Van Horn, D. 1969. Densities of orthopteroid insect popula- of Agriculture, South Dakota State University, Brookings, tions on the Pawnee National Grassland. Joint annual meet- S. Dak. 57006 (hydrology) ing of the Southwestern and Rocky Mountain Division of the J. 0. Harris, Biology Department, Kansas State University, American Association for the Advancemit of Science and Manhattan, Kans. 66502 (soil microbiology) the Colorado-Wyoming Academy of Science, Colorado H. F. Heady, School of Forestry and Conservation, University Springs, Colo. May 7-10. of California, Berkeley, Calif. 94720 (herbage dynamics) Weber, P., and J. Ives. 1970. Studies on alpine grassland com- S. J. Herrmann, Biology Department, Southern ColoradoState munities. U.S. Grassland Biome Seminar, Univ. Colorado. College, Pueblo, Colo. 81005 (phytoplankton) Wesley, D. E., K. L. Knox, and J. G. Nagy. 1969. Water kinetics R. S. Hoffmann, Museum of Natural History, University of in pronghorn antelope. Joint annual meeting of the South- Kansas, Lawrence, Kans. 66044 (small mammal survey) western and Rocky Mountain Division of the American E. W. Huddleston, Entomology Department, Texas Technolog- Association for the Advancement of Science and the ical University, Lubbock, Tex. 79409 (insect systematics) Colorado-Wyoming Academy of Science, Colorado Springs, G. K. Hulett, Division of Biological Sciences, Fort Hays Kansas Colo. May 7-10. State College, Hays, Kans. 67601 (herbage dynamics) Wesley, D., and J. G. Nagy. 1969. Studies of antelope metabo- H. L. Hutcheson, Animal Science Department, South Dakota lism. U.S. Grassland Biorne Seminar, Univ. Wyoming. State University, Brookings, S. Dak. 57006 (herbage dynamics) D. N. Hyder, Agricultural Research Service, U.S. Department of Agriculture, Colorado State University, Fort Collins, Colo. 80521 (large mammals) D. A. Jameson, Natural Resource Ecology Laboratory, Colorado APPENDIX C: PARTICIPANTS State University, Fort Collins, Colo. 80521 (Pawnee Site Director) G. M. Van Dyne, Natural Resource Ecology Laboratory, J. K. Jones, Jr., Museum of Natural History, Department of Colorado State University, Fort Collins, Colo. 805 21 (Biome Systematics, University of Kansas, Lawrence, Kans. 66044 Director; modeling) (small mammal survey) P. H. Baldwin, Zoology Department, Colorado State University, W. M. Klein, Botany Department, Colorado State University, Fort Collins, Colo. 80521 (bird studies) Fort Collins, Colo. 80521 (plant identification) R. E. Bement, Agricultural Research Service, U.S. Department D. P. Knievel, Agronomy Department, University of Wyoming, of Agriculture, Colorado State University, Fort Collins, Laramie, Wyo. 82070 (photosynthesis) Colo. 80521 (large herbivores) D. H. Knight, Botany Department, University of Wyoming, H. D. Blocker, Entomology Department, Kansas State Univer- Laramie, Wyo. 8207C (vegetation structure) sity, Manhattan, Kans. 66502 (insect studies) K. L. Knox, Animal Science Department, Colorado State Uni- R. D. Burman, Agricultural Engineering Section, University of versity, Fort Collins, Colo. 80521 (large mammals) Wyoming, Laramie, Wyo. 82070 (microclimatology) J. Koranda, Lawrence Radiation Laboratory, Livermore, Calif. Martha Christensen, Botany Department, University of Wyoming, 94550 (herbage dynamics) Laramie, Wyo. 82070 (soil microbiology) C. L. Kucera, Botany Department, University of Missouri, Francis E. Clark, Agricultural Research Service, U.S. Department Columbia, Mo. 65201 (biomass-energy relationship) of Agriculture, Colorado State University, Fort Collins, Colo. J. W. Lavelle, Biology Department, Southern Colorado State 80521 (soil microbiology) College, Pueblo, Colo. 81005 (phytoplank ton) V. Cole, Colorado State University, Fort Collins, Colo. 80521 R. J. Lavigne, Plant Science Division, University of Wyoming, (soil phosphorus) Laramie, Wyo. 82070 (insect studies) D. Collins, Botany-Microbiology Department, Montana State J. K. Lewis, Animal Science Department, South Dakota State University, Bozeman, Mont. 5 975 1 (herbage dynamics) iver sity , Brookings, S. Dak. 57006 (herbage dynamics) J. Crabb, Soil Conservation Service, U.S. Department of Agri- J. E. Lloyd; Plant Science Division, University of Wyoming, culture, University of Northern Colorado, Greeley, Colo. Laiamie, Wyo. 82070 (invertebrates) 80631 (mapping) J. V. Mayeux, Microbiology Department, Colorado State Uni- D. A. Crossley, Jr., Entomology Department, University of versity, Fort Collins, Colo. 80521 (soil microbiology) Georgia, Athens, Ga. 30601 (invertebrates) B. McDaniel, Entomology Department, South Dakota State K. G. Doxtader, Agronomy Department, Colorado State Uni- University, Brookings, S. Dak. 57006 (soil arthropods) versity, Fort Collins, Colo. 80521 (soil microbiology) L. D. Miller, Recreation and Watersheds Resources, Colorado G. Ettershank, Department of Biology, Texas Technological State University, Fort Collins, Colo. 80521 (remote sensing) University, Lubbock, Tex. 79409 (soil fauna) W. H. Moir, Range Science Department, Colorado State Univer- H. G. Fisser, Plant Science Division, University of Wyoming, sity, Fort Collins, Colo. 805 21 (photosynthesis) Laramie, Wyo. 82070 (vegetation structure) M. S. Morris, School of Forestry, University of Montana, R. C. Francis, Natural Resource Ecology Laboratory, Colorado Missoula, Mont. 59801 (herbage dynamics)

18 J. G. Nagy, Fishery and Wildlife B:ology Departmen t, Colorado DECIDUOUS FOREST BIOME State University, Fort Collins, Colo. 80521 (large mammals) C. O'Brien, Entomology Department, Texas Technological Uni- versity, Lubbock, Tex. 79409 (insect studies) R. L. Packard, Department of Biology, Texas Technological Deciduous forests are the dominant vegetation in University, Lubbock, Tex. 79409 (small mammal survey) nearly 30 states in the eastern and midwestern United R. M. Pengra, Bacteriology Department, South Dakota State University, Brookings, S. Dak. 57006 (soil microbiology) States, covering about 350 million acres. This acreage is R. D. Pettit, Department of Range Management, Texas Techno- about 52 percent of the total acreage in the two regions logical University, Lubbock, Tex. 79409 (herbage dynamics) and about 50 petcent of the total forest acreage in the R. E. Pfadt, Plant Science Division, University of Wyoming, United States. Laramie, Wyo. 82070 (insect studies) R. D. Pieper, Animal Range and Wildlife Sciences, New Mexico The deciduous forests of the United States are the State University, Las Cruces, N. Mex. 88001 (herbage dy- basis of important lumber and wood-product industries, namics) and they are a valuable recreational and conservation re- L. Pochop, Plant Science Division, University of Wyoming, Laramie, Wyo. 82070 (meteorology) source. The forested cover affects the yield and quality L. K. Porter, Agricultural Research Service, U.S. Department of fresh water from the watershed. of Agriculture, Colorado State University, Fort Collins, The fertile floodplains of drainage basins are among Colo. 80521 (soil nitrogen) J. L. Rasmussen, Atmospheric Science Department, Colorado the most productive agricultural lands 'In the world, and State University, Fort Collin., Colo. 80521 (climatology) most of the nation's inland urban complexes have devel- F. Rauzi, Plant Science Division, University of Wyoming, oped adjacent to these centers of food production. The Laramie, Wyo. 82070 (hydrology) J. 0. Reuss, Agronomy Department, Colorado State University, combination of land and water resources (food, fiber, Fort Collins, Colo. 80521 (soil nitrogen) transportation, and water) supports intensive industrial R. W. Rice, Animal Science Department, University of Wyoming, development and population concentrations. About 75 Laramie, Wyo. 82070 (large mammals) P. G. Risser, Botany Department, University of Oklahoma, percent of the U.S. population lives in the temperate de- Norman, Okla. 37069 (vegetation structure) ciduous forest biome; more than 70 percent of these peo- R. A. Ryder, Fishery and Wildlife Biology Department, Colorado ple live in urban areas. State University, Fort Collins, Colo. 80521 (bird studies) J. A. Seilheimer, Biology Department, Southern Colorado State The deciduous forest biome program is one of six College, Pueblo, Colo. 81005 (zooplankton) biome programs designed to analyze the ecological and S. Shushan, Biology Department, University of Colorado, related physical processes that serve to regulate forested, Boulder, Colo. 80302 (reference collection) P. L. Sims, Range Science Department, Colorado State University, agricultural, and aquatic ecosystems. Through modern Fort Collins, Colo. 80521 (herbage dynamics) integrative and synthesis techniques (e. g., systems anal- J. Smhh, Recreation and Watershed Resources, Colorado State University, Fort Collins, Colo. 80521 (remote sensing) ysis and computer simulation modeling), predictive mod- E. E. Staffeldt, Biology Department, New Mexico State Univers- els of key processes in ecosystems as well as models of ity, Las Cruces, N. Mex. 88001 (soil microbiology) entire ecosystems are being developed. Particular empha- C. L. Streeter, Animal Science Department, Colorado State Uni- versity, Fort Collins, Colo. 80521 (large mammals) sis is being placed on the development of integrated mod- W. D. Striffler, Recreation and Watershed Resources, Colorado els of interacting land and water systems since these sys- State University, Fort Collins, Colo. 80521 (hydrology) tems are being subjected to the greatest stresses. By com- G. Swartzman, Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, Colo. 80521 (data analysis; bining studies of basic ecosystem processes with studies modeling) of landwater systems now undergoing modification by T. 0. Thatcher, Entomology Department, Colorado State Uni- man, it may be possible to derive a scientific basis for re- versity, Fort Collins, Colo. 80521 (insect studies) B. 0. Thomas, Biological Sciences Department, University of source management and for long-term utilization of land Northern Colorado, Greeley, Colo. 80631 (aquatic studies) and water. G. W. Tomanek, Division of Biological Sciences, For t Hays Kansas State College, Hays, Kans. 67601 (herbage dynamics) J. Trappe, Forest Service, U.S. Department of Agriculture, Oregon State University, Corvallis, Oreg. 97330 (aquatic studies) D. H. Van Horn, Biology Department, University of Colorado, RESEARCH PLANS Colorado Springs, Colo. 80907 (insect studies) C. R. Ward, Entomology Department, Texas Technological Uni- versity, Lubbock, Tex. 79409 (aquatic insects) T. Weaver, Botany Department, Montana State University, Research in the deciduous forest biome is planned to Bozeman, Mont. 59715 (herbage dynamics) develop an understanding of forest ecosystems in several F. W. Whicker, RadiLlogy Department, Colorado State Univer- geographic areas and at several 'levels of complexity so sity, Fort Collins, Colo. 80521 (soil dynamics) W. C. Whitman, Department of Botany, North Dakota State that realistic predictions can be made in terms of geo- University, Fargo, N. Dak. 58102 (herbage dynamics) graphical and biological patterns. To achieve this general- J. A. Wiens, Zoology Department, Oregon State University, ity, the program is organized around a number of major Corvallis, Oreg. 97331 (bird survey) sites with representative local and regional characteristics. Studies at these major sites are being augmented

19 by a comprehensive network of individual studies toprovide the necessary geographic coverage. Much ol the program is oriented toward analysis of some of the basic production processes that regulate forest and aquatic ecosystems: biological productivity, nutrient cycling, food chain mechanisms, energy flow, hydrologic cycles, and exchanges between systems. In related studies, ecosystem components will be monitored and changes (e. g., in standing crops of vegetation or in animal biomass) will be monitored. The resulting data will be combined with data from projects in which emphasis was placed on experimental manipulations of ecosystem units. One phase of the experimental treatment will include addition of nutrients (or deletion, as in forest cutting or water treatment techniques) to provide more exact control of variables as a means of determining the extent to which manipulation of one ecosystem component influences the structure and function of other FIGURE 4Sites for research on the deciduous forest components. biome. Although the research will be concerned primarily with the chemical relationships between forest and aquatic ecosystems, the importance of agriculture in this Branch watershed empty into Melton Hill Lake, an em- biome will not be overlooked. Much of the agricultural bayment of the Tennessee River system. The ecology and urban-suburban areas utilize what was originally for- group at the Laboratory is dedicated to integrated eco- est land. Because of future needs for food and fiber, it s) stem research and is one of the largest groups of its seems desirable to study both the quality and the quan- kind in the country. tity of total production in some forest and agricultural 2. Lake Wingra (University of Wisconsin); located at ecosystems. However, these investigations will be post- Madison, Wisconsin. This site offers great potential for poned until the initial research has delineated suitable studying the impact of urban, suburban, and recreational methods and appropriate aspects for emphasis. Social use on a body of water. Research will include the ecology criteria, ecological indices, and system modeling will be of lake basins, hydrology, and related physical systems. combined to design studies and experiments that will The University of Wisconsin group has done distinguished yield widely applicable information. work in limnology and related studies. Systems analysis, which provides the mechanism for 3. Lake George (Rensselaer Polytechnic Institute); investigators from many different disciplines to work to- located on the eastern edge of the Adirondack province. gether, will operate at all levels of research planning, data Lake George is an oligotrophic lake that is receiving in- analysis, and model synthesis. creased amounts of pollution from the resort development in the southem subbasin. Groups at RPI and several branches of the State University of New York have SITES organized an enthusiastic team of investigators. 4. Coweeta (University of Georgia and U.S. Forest Five major sites have been selected for process studies Service); located in the basin of the upper Tennessee in an ecosystem context (Figure 4). They provide a com- River. The stream system is on base-poor crystalline bed- bination of environmental situations typical of bnajor rock, and studies of interaction between land and water parts of the biome. By grouping process studies at each are being conducted here. Data from these studies will be of these sites, it will be possible to assemble and organize compared with those from studies in the lime-rich basin ecosystem analyses at different levels of complexity and of the Oak Ridge site and with those from the Hubbard for different geographical coverage. Brook (New Hampshire) study. The University of Georgia 1. Oak Ridge (Oak Ridge National Laboratory); lo- is the center of a well-known ecology group. cated in the basin of the upper Tennessee River. This site 5. Triangle (U.S. Forest Service, Duke University, has a major facility for studying interactions between University of North Carolina, and North Carolina State land and water. Water and related terrestrial nutrient cy- University); located on the edge of the Piedmont Plateau cling is being investigated in dolomitic limestone terrain. near the Costal Plain. This site is characterized by second- The streams draining the well-instrumented Walker ary southern pine forests normally replaced by deciduous

22 forest. Work here emphasizes photosynthesis, forest pro- related (e. g., primary productivity to secondary produc- ductivity, and related meteorology. The Triangle group tion to decomposition). The integration of these sub- has a large number of ecologists, and two major phy- system processes is the responsibility of the site coordi- totrons are available, one at Duke University and the nators. Finally, the overall integration of results and other at North Carolina State University. development of biome models is the responsibility of the Selected process studies are being conducted at other Biome Director. sites by other groups. The additional studies are analo- gous to the "comprehensive" programs of the other biomes, being intended to augment the data base. Ini- tially, only a few are being funded from the grant for the RESEARCH METHODS AND PROGRESS deciduous forest biome. Inclusion of more sites will depend on the development and progress of research at the five major sites. Initial funding for the biome was made available in 1969. It provided for initiation of modeling research and lead-time investigations on certain process studies. Model- ORGANIZATION ing research was begun at the Oak Ridge, Lake Wingra, Lake George, and Triangle sites and in the Biome Di- S. I. Auerbach is the Deciduous Forest Biome Di- rector's office. Initial studies of water chemistry and rector. He is assisted by R. L. Burgess, who is Deputy aquatic primary productivity were started at Lake Wingra Director, by an administrative assistant, and by research and Lake George; work on primary production in terres- coordinators. The organizational structure below the trial ecosystems was started at the Triangle site. level of the Deputy Director and the administrative as- Thc: modeling program at the Lake Wingra site has sistant is shown in the following list: focused on the movement of energy and carbon in the aquatic system and the coupling of biomass simulation A. International coordination (J. S. Olson) capability with nutrient and water environment simula- B. Interbiome coordination (F. Glenn Goff) tions. Energy-transfer, water-transfer, and nitrogen- C. Resource region modeling and site modeling transfer flow charts have been developed in matrix form. I. Coordinator (R. V. O'Neill) The energy output at each juncture of the matrix involves 2. Modeling committee a process: reflection, absorption, photosynthesis, reradia- D. Major sites tion, vaporization, condensation, respiration, fire, trans- I. Oak Ridge (D. E. Reichle, coordinator) port (migration, litter fall), ingestion (grazing, predation), 2. Lake Wingra (0. Loucks, coordinator) assimilation, excretion, decomposition, mortality, and 3. Lake George (N. Clesceri, coordinator) replication. These studies have contributed to the develop- 4. Coweeta (C. D. Monk, coordinator) ment of a more detailed research design for the studies 5. Triangle (B. Strain, coordinator) land and water interaction. E. Process studies Modeling of the Lake George aquatic ecosystem uses I. Primary productivity empirical relationships among diatoms, key nutrients, 2. Secondary productivity and other chemical and physical factors. The construc- 3. Decomposition processes and soils tion and evaluation of certain models have been aided by 4. Land-water interactions regression analyses and by graphical procedures where 5. Streams the data were sparse or incomplete. These analyses in- 6. Lakes dicated (1) that both phosphorus and nitrogen are limit- F. Comprehensive and related studies ing at certain times, and other nutrients and growth factors may be limiting; (2) that nutrient levels vary sea- Technical modeling services, data processing, editorial sonally (apparently in response to seasonal influxes of services, chemical services, and remote sensing services vacationers); and (3) that nutrient and corresponding are provided by the central biome office. productivity gradients are closely related to summer pop- The first level of synthesis is with the research teams ulation density. working on ecosystem processes. These teams synthesize Simple compartment models, based partly on the ob- data into relevant forms, developing submodels of the served relationships and partly on published material, processes and working toward general process models. A have been designed and programmed. From these initial second level of integration takes place at each site. Here models, it is evident that the relationships between the the different process studies are brought together so that primary producer compartment and the available nutrient the data for modeling and simulation are appropriately compartment are crucial. Because several nutrients may

23 be at threshold levels at a given time, relatively detailed Robert L. Barnes, School of Forestry, Duke University, Durham, modeling is required. Programming is being initiated to N.C. (forest biochemistry) E. W. Beals, Department of Zoology, University of Wisconsin, develop models that can be easily modified as more ad- Madison (avian ecology) equate empirical models are developed for the compart- Donald Bishko, Industrial Administration and Computer Science, ment components. Union College, Schenectady, N.Y. (computer science) James Reid Boyle, Department of Soil Science, University of At the Triangle site, a group at Duke University has Wisconsin, Madison (soil science) been modeling a deciduous forest stand energy budget. Alfred A. Brooks, Jr., Head, Scientific Applications Department, The energy exchange between a forest stand and the en- Computing Technology Center, Oak Ridge Gaseous Diffusion Plant, Oak Ridge, Tenn. (computer science) vironment surrounding it has been simulated by finding a Stanley W. Buol, Soil Science Department, North Carolina State simultaneous solution to a set of equations (based on University, Raleigh (soil science) continuum thermodynamics and fluid mechanics) that Edmondo Canelli, Division of Laboratories and Research, New York State Department of Health, Albany, N.Y. (environ- describe the energy exchange process. While this model mental microbiology) does not directly estimate the photosynthesis or res- Min Chen, Division of Laboratories and Research, New York piration of forest stands, it does provide a quantitative State Departmcnt of Health, Albany, N.Y. (environmental microbiology) description of the stand environment that can be used to James L. Clapp, Civil Engineering Department, University of make indirect estimates of these quantities. Wisconsin, Madison (civil engineering) A technical modeling group has been formed at the Edward E. C. Clebsch, Department of Botany, University of Tennessee, Knoxville (plant ecology) Oak Ridge site. This group is charged with providing tech- Lenore S. Clesceri, Department of Biology, Rensselaer Polytech- nical support to the modeling effort throughout the nic Institute, Troy, N.Y. (aquatic biochemistry) biome and with developing a general forest ecosystem Nicholas L. Clesceri, Environmental Engineering, Rensselaer Polytechnic Institute, Troy, N.Y. (site coordinator) model. It has encouraged publication of a data retrieval David C. Coleman, Department of Zoology, University of program specifically designed for ecological data. Other Georgia, Athens (soil ecology) tasks in progress include an annotated bibliography of Arthur W. Cooper, Department of Botany, North Carolina State University, Raleigh (biomass determination) modeling literature, a review of relevant computer tech- Grant Cottam, Department of Botany, University of Wisconsin, niques and programs, and a study on methods of solving Madison (plant ecology) differential equations in ecosystem models. D. A. Crossley, Entomology Department, University of Georgia, Athens (insect and soil ecology) J. W. Curlin, Ecological Sciences Division, Oak Ridge National Laboratory, Oak Ridge, Tenn. (fOrest ecology and hydrology) Roger C. Dahlman, Ecological Sciences Division, Oak Ridge National Laboratory, Oak Ridge, Tenn. (plant ecology; nitro- APPENDIX A: PUBLICATIONS gen cycling) Robert J. Dicke, Entomology Department, University of Wiscon- Baskerville, G. L. 1970. Testing the uniformity of variance in sin, Madison (entomology) arithmetic and logarithmic units of a Y-variable for classes Frederick Dreher, Water Resources Division, Geological Survey, of an X-variable. ORNL-IBP-60-1. (Oak Ridge National U.S. Department of the Interior, Madison, Wis. (geology) Laboratory, Tenn.) Paul B. Dunaway, Ecological Sciences Division, Oak Ridge Hull, Norma, J. Beauchamp, and D. Reichle. 1970. Pitfall I: a National Laboratory, Oak Ridge, Tenn. (vertebrate ecology) general purpose data processing program for environmental Nelson T. Edwards, Ecological Sciences Division, Oak Ridge data. ORNL-IBP-70-2. (Oak Ridge National Laboratory, National Laboratory, Oak Ridge, Tenn. (decomposition Tenn.) processes) Reichle, D. [ed.] 1970. Analysis of temperate forest ecosystems. Alan R. Ek, Department of Forestry, University of Wisconsin, (Based on IBP/PT meeting held in Gatlinburg, Tenn.) Madison (forest ecology) Springer-Verlag, Heidelberg and New York. Jerry W. Elwood, Ecological Sciences Division, Oak Ridge Na- Taylor, F. 1969. Phenological records of vascular plants at Oak tional Laboratory, Oak Ridge, Tenn. (aquatic ecology; secon- Ridge, Tennessee. ORNL-IBP-69-1. (Oak Ridge National dary productivity) Laboratory, Tenn.) Jerald C. Ensign, Department of Bacteriology, University of Wisconsin, Madison (microbiology) Charles H. Falkner, Industrial Engineering Department, Univer- sity of Wisconsin, Madison (computer and systems science) John M. Ferris, Entomology Department, Purdue University, APPENDIX B: PARTICIPANTS Lafayette, Ind. (entomology) George P. Fitzgerald, Department of Water Chemistry, University Stanley I. Auerbach, Director, Ecological Sciences Division, Oak of Wisconsin, Madison (water chemistry) Ridge National Laboratory, Oak Ridge, Tenn. (Biome Chester W. Francis, Health Physics Division, Oak Ridge National Director) Laboratory, Oak Ridge, Tenn. (soils) Michael S. Adams, Department of Botany, University of Wiscon- Gerald M. Friedman, Department of Ecology, Rensselaer Poly- sin, Madison (aquatic primary productivity) technic Institute, Troy, N.Y. (geology) D. E. Armstrong, Department of Water Chemistry, University of G. Wolfgang Fuhs, Environmental Health Center, Division of Wisconsin, Madison (water chemistry) Laboratories and Research, New York State Department of Richard Warren Arnold, Department of Soil Science, Cornell Health, Albany, N.Y. (aquatic microbiology) University Agricultural Experiment Station, Ithaca, N.Y. Wilford R. Gardner, Soils Department, University of Wisconsin, (soils) Madison (soil and water physics) Allen Baisuck, Rensselaer Research Corporation, Troy, N.Y. R. L. Giese, Department of Entomology, Purdue University, (operations analysis) Lafayette, Ind. (insect ecology)

22 24 F. Glenn Goff, Ecological Sciences Division, Oak Ridge National Edward E. Miller, Department of Physics and Soils, University Laboratory, Oak Ridge, Tenn. (forest ecology) of Wisconsin, Madison (physics and soils) Robert A. Goldstein, Ecological Sciences Division, Oak Ridge John W. Mitchell, Department of Mechanical Engineering, Uni- National Laboratory, Oak Ridge, Tenn. (computer and versity of Wisconsin, Madison (systems engineering) systems science) Carl D. Monk, Department of Botany, University of Georgia, Joseph B. Gonthier, Geological Survey, U.S. Department of the Athens (site coordinator) Interior, Madison, Wis. (geology) Peter L. Monkmeyer, Department of Civil Engineering, Univer- Theodore Green, III, Department of Civil Engineering and Mete- sity of Wisconsin, Madison (civil engineering) orology, University of Wisconsin, Madison (civil engineering; Charles E. Murphy, Duke University, Durham, N.C. (energy meteorology) exchange) Robin F. Harris, Department of Soils, University of Wisconsin, Daniel J. Nelson, Ecological Sciences Division, Oak Ridge Na- Madison (soils) tional Laboratory, Oak Ridge, Tenn. (aquatic ecology) W. Franklin Harris, Ecological Sciences Division, Oak Ridge Robert V. O'Neill, Ecological Sciences Division, Oak Ridge National Laboratory, Oak Ridge, Tenn. (primary production) National Laboratory, Oak Ridge, Tenn. (systems ecology) Arthur D. Hasler, Director, Laboratory of Limnology, University R ichard A. Park, Department of Geology, Rensselaer Polytechnic of Wisconsin, Madison (aquatic ecology) Institute, Troy, N.Y. (systems analysis) Henry Hellmers, Department of Botany, Duke University, Bernard C. Patten, Department of Zoology, University of Durham, N.C. (plant physiology) Georgia, Athens (systems ecology) Joan M. Hett, Ecological Sciences Division, Oak Ridge National Warren P. Porter, Department of Ecology, University of Wis- Laboratory, Oak Ridge, Tenn. (forest ecology) consin, Madison (physiological ecology) William L. Hilsenhoff, Department of Entomology, University Robert A. Ragotzkie, Department of Meteorology, University of Wisconsin, Madison (aquatic entomology) of Wisconsin, Madison (meteorology) Francis D. Hole, Department of Soil Science, University of Charles W. Ralston, Dean, School of Forestry, Duke University, Wisconsin, Madison (soils) Durham, N.C. (forest soils) Lee Holt, Water Resources Division, Geological Survey, U.S. William G. Reeder, Department of Zoology, University of Wis- Department of the Interior, Madison, Wis. (geology) consin, Madison (vertebrate ecology) J. Gary Holway, Department of Biology, State University David E. Reichle, Ecological Sciences Division, Oak Ridge Na- College, Oneonta, N.Y. (plant ecology) tional Laboratory, Oak Ridge, Tenn. (site coordinator) John A. Hoopes, Department of Civil Engineering, University Orrin Rongstad, Wildlife Extension Specialist, University of of Wisconsin, Madison (environmental engineering) Wisconsin, Madison (wildlife ecology) Dale D. Huff, Department of Civil Engineering, University of Michael L. Rosenzweig, Department of Biological Sciences, Wisconsin, Madison (hydrology) State University of New York, Albany, N.Y. (ecology) Kermit Hutcheson, Department of Statistics, University of John E. Ross, Department of Agricultural Journalism, University Georgia, Athens (statistics) of Wisccnsin, Madison (agricultural journalism) James H. Jenkins, School of Forestry, University of Georgia, Manfred Schoettle, Department of Geology, Rensselaer Poly- Athens (wildlife ecology) technic Institute, Troy, N.Y. (geology) Jacques R. Jorgensen, Forestry Sciences Laboratory, U.S. Jon T. Scott, Atmospheric Sciences Research Center, State Uni- Forest Service, Research Triangle Park, N.C. (forest soils) versity of New York, Albany (atmospheric sciences) D. R. Keeney, Department of Soil Science, University of Earl L. Skinner, Water Resources Division, Geological Survey, Wisconsin, Madison (soils) U.S. Department of the Interior, Madison, Wis. (hydrology) Kenneth R. Knoerr, Department of Forest Meteorology and Nancy F. Sollins, Ecological Sciences Division, Oak Ridge Na- Biometeorology, Duke University, Durham, N.C. (photo- tional Laboratory, Oak Ridge, Tenn. (information and data synthesis) processing) Edward J. LaRow, Department of Biology, Siena College, Philip Sollins, Department of Botany, University of Tennessee, Loudonville, N.Y. (aquatic ecology) Knoxville (systems ecology) Gerhard B. Lee, Department of Soil Science, University of Richard Duane Spear, Division of Bioenvironmental Engineering, Wisconsin, Madison (soils) Rensselaer Polytechnic Institute, Troy, N.Y. (bioenvh on- G. Fred Lee, Department of Water Chemistry, University of mental engineering) Wisconsin, Madison (water chemistry) C. R. Stearns, Division of Meteorology, University of Wisconsin, Bernhard Lettau, Department of Meteorology, State University Madison (meteorology) of New York, Albany, N.Y. (meteorology) Ronald Stewart, Department of Atmospheric Sciences, State Philip H. Lewis, Jr., Department of Landscape Architecture, University of New York, Albany (atmospheric sciences) University of Wisconsin, Madison (landscape architecture) Boyd R. Strain, Department of Botany, Duke University, Helmut F. Lieth, Department of Botany, University of North Durham, N.C. (photosynthesis) Carolina, Chapel Hill, N.C. (primary productivity) Raymond G. Stross, Department of Biology, State University Orie L. Loucks, Department of Botany, University of Wisconsin, of New York, Albany (aquatic ecology) Madison (site coordinator) J. K. Syers, Department of Soils, University of Wisconsin, John J. Magnuson, Department of Zoology, University of Wis- Madison (soils) consin, Madison (fisheries ecology) T. Tamura, Health Physics Division, Oak Ridge National Labora- James B. Mankin, Jr., Computing Technology Center, Oak Ridge tory, Oak Ridge, Tenn. (soils) Gaseous Diffusion Plant, Oak Ridge, Tenn. (cow puter sci- Champ B. Tanner, Department of Soila, University of Wisconsin, ence) Madison (soils) Michael D. McCracken, Department of Botany, University of Fred G. Taylor, Jr., Ecological Sciences Division, Oak Ridge Wisconsin, Madison (botany) National Laboratory, Oak Ridge, Tenn. (forest ecology) Robert S. McLeod, Geological Survey, U.S. Department of the Gerald U. Uhickson, Ecological Sciences Division, Oak Ridge Interior, Madison, Wis. (hydrology) National Laboratory, Oak Ridge, Tenn. (aquatic ecology; Donald C. McNaught, Department of Biological Sciences, State information storage and retrieval) University of New York, Albany, N.Y. (aquatic ecolly; J. Bruce Wallace, Department of Entomology, University of secondary productivity) Georgia, Athens (entomology) Louis J. Metz, Soil Scientist, U.S. Forest Service, Research Tri- William A. Wallace, School of Management, Rensselaer Poly- angle Park, N.C. (forest soils) technic Institute, Troy, N.Y. (computer science) Clarence J. Milfred, Soil Survey Division, University of Wisconsin, Donald G. Watts, Statistics Department, University of Wiscon- Madison (soils) sin, Madison (statistics)

23 25 Carol G. Wells, Southeastern Forest Experiment Station, Re- During the initial phase of this program extensive use search Trianglc Park, N.C. (soil-tree relations) is made of past studies. Existing models of biomass, min- William F. Whittingham, Department of Botany, University of Wisconsin, Madison (soil microbiology) eral cycling, primary production, and population dynam- Richard G. Wiegert, Department of Zoology, University of ics that exist for a number of ecosystems at selected Georgia, Athens (invertebrate ecology) intensive study sites are being further analyzed, tested, John W. Wilkinson, School of Management, Rensselaer Poly- technic Institute, Troy, N.Y. (operations research; statistics) and applied to new communities. New field studies will Cliff R. Willey, Biological and Agricultural Engineering, North be initiated as they are suggested by the initial modeling Carolina State University, Raleigh (soil physics) John P. Witherspoon, Ecological Sciences Division, Oak Ridge program. In these new studies, careful attention will be National Laboratory, Oak Ridge, Tenn. (forest ecology) given to relations between the aquatic and terrestrial Martin Witkamp, Ecological Sciences Division, Oak Ridge Na- components. tional Laboratory, Oak Ridge, Tenn. (microbial ecology) David J. Yount, Institute of Ecology, Athens, Ga. (chemistry) Later phases will emphasize: James H. Zimmerman, Phenology Department, University of Expansion of current studies to the other important Wisconsin Aboretum, Madison (phenology) ecosystems Continued development of the process and validation studies in ways suggested by the modeling programs and simulation studies Coordination of existing process studies and the CONIFEROUS FOREST BIOME modeling programs throughout the biome Development of new biome-wide studies that will help in understanding basic processes and behavior within Because of diversity of environments, species, soils, the biome products, and land uses, single af ,>roaches to managing We are not attempting to construct biome-wide simu- the western coniferous forest are impossible. The use and lation models during early stages of the work. Instead, misuse of the forest by modern colonizers moving west- investigators and modelers at each site will develop models ward led to complicated efforts to understand the forest. of local subsystems and combine these with computer The time-honored use of the forest as a source of agricul- models of the local ecosystems. Frequent seminars, con- tural land is being replaced by use for suburban and rec- ferences, and visits enable modelers at the intensive and reational development. As a result, large numbers of peo- coordinating sites to maintain close contact with one ple are impinging on sensitive ecosystems. another and with modelers working on other biomes. In the final stages, emphasis will be placed on integration of research results. Biome-wide models of processes in subsystems, such as models describing movement of RESEARCH PLANS water and nutrients through soils or the growth and development of certain organisms that are widely distributed over the biome, will be attempted. In addition, models of Research in the coniferous forest biome will include entire local ecosystems will be compared and the feasibil- process studies, computer modeling, and validation ity of constructing biome-wide models for whole eco- studies, in that sequence. As discrepancies are encoun- systems will be studied. tered between the predictions of the simulation model and field observations and validation studies, explanations will be sought and new process studies will be sug- SITES gested. The new studies will refine and extend the range of applicability of the functions that will be incorporated The coniferous forest biome encompasses a wide range in local, and eventually biome-wide, ecosystem models. of forest ecosystems varying markedly in environment, Since our purpose is to increase understanding of composition, structure, and productivity. These eco- whole ecosystems within the coniferous forest biome, we systems are managed differently because their resources study processes and develop models that are relevant to are valued differently. A general model of coniferous for- entire systems. Accordingly, each project, however nar- est ecosystems must reflect reality by encompassing this row or detailed, will develop a submodel that relates to variability in intrinsic character and manipulative prac- an essential component of the system. At all major sites tice. For this reason, research will be carried out at within the biome, simulation models are being developed several study sites. Further, a requirement for data relat- that describe the behavior of subsystems within the ing one process to another quantitatively and for linking local ecosystem. Later, simulation models for entire local the terrestrial and aquatic components of the ecosystems ecosystems will be developed by combining models of makes it essential that representative sites have a history local subsystems. of research development and sufficient scientific man-

24 26 The drainage basin of Lake Washington consists of two distinct subdrainages, the Sammamish and Cedar River valleys in central Washington. These two drainage systems differ greatly in both physiography and the patterns of land use. The Lake Sammamish system, which includes Lake Sammamish, the Sammamish Slough, and a variety of tributary systems, including Issaquah Creek, has relatively low elevation and is subject to heavy pressures of urbanization and agricultural practices. Although a series of restrictions have been placed on direct pollution of these water systems, the area still receives a very high input of nutrients through secondary sources. The area drained by the Cedar River system has an elevation higher than that drained by the Lake Sammamish system, and it consists primarily of forest ecosystems. The Cedar River system includes several small lakes at an elevation of more than 3,500 ft, a large reservoir system at an elevation of 1,555 ft, and Cedar River. The parts of the system that have an elevation of more than 600 feet belong to the watershed from which the city of Seattle receives its water supply. Consequently, this area is carefully protected, and access is rigidly controlled. The forest ecosystems have been systematically harvested and FIGURE 5Sites for research on the coniferous forest reforested during the past 60 years, and the proce.r...z has biome. provided a wide range of forest ages and types. amate is maritime. Precipitation ranges between 50 and ltIO power to conduct the research. Fortunately, several such inches, the amount depending on elevation. Se-0.s are di- sites are available (Figure 5). verse in nature and origin. The soils at lower t.1...tvc:tions The initial biome research is carried out on two inten- range from alluvial to glacial outwash. The soils at higher sive sites: the H. J. Andrews Experimental Forest and the elevations consist of residuum from the weathering of drainage basin of Lake Washington. For both areas con- Andesite with a volcanic ash cover. siderable inventory and process data are already available Nine coordination sites have been proposed. Their and will form the basis for initial modeling activity, along names and the states in which they are located are as fol- with data from Castle Lake in California and Wood River lows: Cache la Poudre River, Colorado; Castle Lake, Cal- Lakes in Alaska. As the research activity expands, addi- ifornia; Fairbanks area, Alaska; Flathead Lake, Montana; tional sites for coordinated validation studies will be Eagle Lake, California; Little South Priest Lake, Idaho; selected. upper Logan Canyon, Utah; White Canyon-Cedar Mesa, The H. J. Andrews Experimental Forest is a 15,000- Utah; Wood River Lakes, Alaska. acre drainage basin in the Oregon part of the Cascade Range and is administered by the Pacific Northwest For- est and Range Experiment Station, U.S. Forest Service. ORGANIZATION The environment and vegetation are typical of much of western Oregon and Washington. Old-growth (450 years Overall policy is developed by the Executive Com- old) Douglas-fir and western hemlock forests are dominant mittee. Stanley P. Gessel is Biome Director. He is assisted in areas below 3,000 ft. Above 3,000 ft true fir and hem- by two Deputy Directors and an administrative assistant. lock forests are dominant; however, extensive areas of The rest of the organizational structure is shown in the Douglas-fir, 60 to 125 years old, and 1,500 acres of cut- following list: overs, up to 16 years old, are also present. Mature topography, elevations of 1,500 to 5,300 ft, volcanic bedrock, A. Planning and Program Design Committee (directors; reddish-brown lateritic and podzolic soils, and a maritime chairmen of research and site area committees) climate with annual rainfall of 94 inches characterize the B. Research committees environment. A market evaluation for the resources 1. Terrestrial produ .ers would rank wood production, water protection, and rec- a. Biomass and structure reational use (camping, fishing, and hunting) among the b. Processes highest management priorities. 2. Terrestrial consumers

25 27 3. Terrestrial decomposers APPENDIX A: PUBLICATIONS 4. Aquatic Ballard, T. M. 1968. Carbon dioxide production and diffusion 5. Aquatic-terrestrial interface in forest floor material: a study of gas exchange in biologi- a. Nutrient cycling cally active, porous media. PhI:. dissertation. Univ. Washing- ton, Seattle. b. Hydrology Cole, D. W., and P. S. Machno. 1969. Factors affecting percola- c. Meteorolcgy tion in forest soiis. In Proceedings of Third Annual Sympo- 6. Genetics sium of the American Water Research Association. Banff, 7. Modeling Alberta, Canada Cole, D. W., and S. F. Dice. In press. Biomass and nutrient flux C. Technical committees in coniferous forest ecosystems: the development of a quan- 1. Remote sensing titative ecological approach. In Proceedings of Sympcsium on 2. Communication Coniferous Forest of the Northern Rocky Mountains. Univ. Montana, Missoula, September 17-20,1968. 3. Chemical analysis Cole, D. W., S. P. Gessel, and S. F. Dice. 1967. Distribution and 4. International cycling of nitrogen, phosphorus, potassium, and calcium in 5. Data bank a second-growth Douglas-fir ecosystem. Symposium on Pri- mary Productivity and Mineral Cycling in Natural Ecosys- D. Site area committees tems. Ecological Society of America, AAAS annual meeting, 1. Intensive sites New York, N.Y. Univ. Maine Press, Orono. a. H. J. Andrews site Dyrness, C. T. 1967. Mass soil movements in the H. J. Andrews Experimental Forest. U.S. Forest Service. Res. Paper PNW-42. b. Drainage basin of Lake Washington 12 p. 2. Coordination sites Franklin, J. F., and C. T. Dyrness. 1969. Vegetation of Oregon and Washington. U.S. Forest Service Res. Paper PNW-80. 216 p. Levno, A., and J. Rothacher. 1967. Increases in maximum stream temperatures after logging in old-growth Douglas-fir watersheds. U.S. Forest Service Res. Note PNW-65.12 p. RESEARCH METHODS AND PROGRESS McColl, J. G., and D. W. Cole. 1968. A mechanism of cation transpori in a forest soil. Northwest Sci. 42: 134-140. Rothacher, J. 1965. Streamflow from small watersheds on the western slope of the Cascade Range cf Oregon. Water Resour. Since the coniferous biome subprogram was not Res. 1:125-135. funded until September 1970, research results have been Rothacher, J., C. T. Dyrness, and R. L. Fredriksen. 1967. Hy- limited. drologic and related properties of three small watersheds in the Oregon Cascades. U.S. Forest Service, Pacific Northwest Results of studies on mineral cycling and biomass at Forest and Range Experiment Station. 54 p. Cedar River are well documented, and various publications describe the characteristics of the principal watersheds at H. J. Andrews. A summary of this work shows the effect of certain forest practices on the quantity and APPENDIX B: PARTICIPANTS quality of water in experimental streams. A unique development has been the perfection lf a proportional Stanley P. Gessel, College of Forest Resources, University of Washington, Seattle (Biome Director) automatic water sampling device for use in studies of Bruce Bare, College of Forest Resources, University of Washing- stream water quality and mineral cycling. Considerable ton, Seattle (Chairman, Technical Committee on Data Bank) information on elemental composition of surface drain- Mark J. Behan, Department of Botany, University of Montana, Missoula (Chairman, Technical Committee on Chemical age waters is now available for mineral cycling models. Analysis) The principal types of vegetation have been described George Brown, School of Forestry, Oregon State University, and the soils have been mapped. Scientists planning and Corvallis (Chairman, Interface Research Subcommittee on Hydrology) conducting biome research on the H. J. Andrews Exper- Robert Burgner, College of Fisheries, University of Washington, imental Forest have been using other financial resources. Seattle (member of Executive Committee; Chairman, Re- The results of this research will be used in an initial mod- search Committee on Aquatic Component) Douglas Chapman, College of Forest Resources, University of eling effort. Washington, Seattle (member of Executive Committee; Large quantities of data on the nutrient status and Chairman, Research Committee on Modeling) productivity of Lake Washington are available from past Dale W. Cole, College of Forest Resources, University of Wash- ington, Seattle (member of Executive Committee; Director, and current work. Considerable information has been de- Cedar River-Lake Washington Site) veloped on the soils, on water and elemental movement, Robert Colwell, School of Forestry and Conservation, Univer- on standing biomass, and on the effect of some manipula- sity of California, Berkeley (Chairman, Technical Committee on Remote Sensing) tions (e. g., addition of elements and forest removal). Ted Dyrness, Forest Sciences Laboratory, Oregon State Univer- Considerable information has been developed for sity, Corvallis (Chairman, Research Committee on Terres- Castle Lake, and it will be the basis for modeling research trial Producers) Jerry Franklin, Forest Science Laboratory, Oregon State Univer- in the initial 1:)iome work, similarly information is avail- sity, Corvallis (Deputy Biome Director, Ecology; member of able for sites in California and Alaska. Executive Committee)

26 Richard L. Fredriksen, Forestry Sciences Laboratory, P.O. Box knowledge concerning the ecosystems in this region 887, Corvallis, Oreg. (Chairman, Interface Research Subcom- would have scientific and practical value. mittee on Nutrient Cycling) Leo J. Fritschen, College of Forest Resources, University of The arid lands of North America are divided into four Washington, Seattle (Chairman, Interface Research Subcom- main regions, in each of which desert biome studies are mittee on Meteorology) in progress. The regions are: Arden R. Gaufin, Director, Environmental Biology, University of Montana, Missoula (member of Executive Committee) The Great Basin: cold deserts with continuous James D. Hall, Forest Sciences Laboratory, Oregon State Uni- snow cover in winter, mainly at an altitude above versity, Corvallis (Chairman, Research Committee on Aquatic- 1,000 m, in the region between the Rocky Mountains Terrestrial Interface) Robert Harris, Pacific Northwest Forest Experiment Station, and the ranges along the Pacific coast. The precipitation U.S. Forest Service, Portland, Oreg. (member of Executive is 200 to 300 mm. Committee; Chairman, Technical Committee on Communi- The Mohave Desert: in the interior of California cation) George E. Hart, Forest Services, Utah State University, Logan and southern Nevada, mainly at an altitude of 500 m or (member of Executive Committee) more, with cool winters and very hot summers. The pre- William H. Hatheway, College of Forest Resources, University cipitation is usually under 200 mm and is highly irregular. of Washington, Seattle (Deputy Biome Director, Modeling) Edwin Mogren, College of Forestry, Colorado State University, The Sonoran Desert: in southern Arizona and Fort Collins (member of Executive Committee; Chairman, northwestern Mexico. The precipitation is 200 to coordinating sites) 300 mm; seasonal peaks occur in summer and winter. D. W. Rains, Department of Agronomy and Range Science, University of California, Davis (member of Executive Com- Winters are warm, summers hot. mittee) The Chihuahuan Desert: in New Mexico, western Hans Riekerk, College of Forest Resources, University of Texas, and northeastern Mexico. The precipitation is 200 Washington, Seattle (Administrative Assistant) George Staebler, Forestry Research Center, Weyerhaeuser to 300 mm; a seasonai peak occurs in summer. Company, Centralia, Wash. (member of Executive Com- To understand the biological basis of productivity in mittee) these ecosystems, we must first understand their dynam- Reinhard F. Stettler, College of Forest Resources, University of Washington, Seattle (Chairman, Research Committee on ics and functioning. If this understanding is wide and Genetics) deep enough, it can aid greatly in predicting effects of Ronald W. Stark, Dean and Coordinator of Research, The pressuyes, stresses, and manipulations on the e,;osystems, Graduate Division, University of Idal.o, Moscow (member of Executive Committee) and hence can direct their wise use and management. Richard D. Taber, College of Forest Resources, University of Washington, Seattle (Chairman, Research Committee on Terrestrial Consumers) Frieda Taub, College of Fisheries, University of Washington, Seattle (Chairman, Research Committee on Decomposers) RESEARCH PLANS Keith Van Cleve, Institute of Northern Forestry, University of Alaska, College (member of Executive Committee) Richard B. Walker, Botany Department, University of Washing- ton, Seattle (Chairman, Producers Research Subcommittee on Research activities are concentrated on building simu- Processes) lation models of the desert ecosystems. Process studies Richard Waring, Forest Research Laboratory, Oregon State University, Corvallis (Director, H. J. Andrews site; Chai-man, needed to provide information for model building, and Technical Committee on International Component) validation studies to check the performance of the Donald M. Wootton, Department of Biology, Chico State College, models, are included. Chico, Calif. (member of Executive Committee) Paul Zinke, School of Forestry and Conservation, University of California, Berkeley (member of Executive Committee) VALIDATION STUDIES

In all the validation studies a complete inventory of the biota will be taken initially, abiotic conditions will DESERT BIOME be monitored continuously, and changes in the biota will be recorded at regular intervals to provide data for comparison with the output of the computer model. The desert biome subprogram is concerned with the Ten sites, two of which are for aquatic studies only, shrub-dominated ecosystems in the arid and semiarid have been selected for validation studies (Figure 6). The lands of the western and southwestern United States, sites are well distributed through the desert areas, have where the annual precipitation is less than 300 mm. This sufficient experienced manpower available, are accessible, program also has marginal relevance to those areas where and are free from interference. Of these 13 sites, 6 will the annual precipitation is between 300 and 400 mm. be in operation by 1971. They are: They are exceptionally vulnerable to ill-considered use Curlew Valley, Idaho and Utah; altitude 1,300- and, in many places, are subject to rapid change. Increased 1,600 m. The site lies near the center of the Great Basin.

27 dominated by Cercidium spp., Prosopis juliflora, and Opuntia spp. One of these wiil be subject to shrub clear- ing by chemical treatment, one will be subject to burning, and the other two will be differentiated by permitting or preventing cattle grazing. Two additional areas will be located in the Avra Valley bajada, west of Tucson. In this bajada, Larrea divaricata and Cercidium micro- phyllum share dominance with various cacti. Larrea shrubs will be cleared from one area and the other will be left undisturbed. Jornada, New Mexico; altitude 1,400 m; precipitation 210 mm. An area northeast of Las Cruces represents the Chihuahuan Desert. A well-defined catchment in this area drains into a seasonal playa. Two adjacent areas are included in the site. Shrubs will be cleared from one area and the other will be left undisturbed. Initiation of validation studies at the other sites has been postponed until funding difficulties are resolved. The other sites are: Hanford Reservation, Washington; altitude 200 m; precipitation 180 mm. The site lies nea- the northern limit of the U.S. cold desert. It includes sagebrush vegetation that was cleared 25 years ago and is reverting to its original state. It also includes a creek that will be the site of the aquatic studies. FIGURE 6Sites for validation studies of the desert biome. Alkali Flat, Wyoming; altitude 200 m; precipitation 250 mm. The site lies in the Red Desert of Wyoming. Two areas have been selected; one is dominated by Arte- Two areas have been selected, both of which have sage- misia tridentata and the other by Atriplex gardneri. brush as the native dominant plant. One area is near the Pine Valley, Utah; altitude 1,600 m; precipitation upper rainfall limit and the other is near the lower rain- 150 mm. The site is near the southern edge of the Great fall limit of the valley (200 and 350 mm). Each area is Basin. Four areas of vegetation, dominated by Atriplex matched with a similar one that is subjected to reseeding. confertifolia, will Le subject to different kinds of grazing Three aquatic studies are also to be conducted, one in a management. small reservoir, one in a creek, and one in a saline spring. Deep Canyon, California; altitude 300-1,000 m; Saratoga Springs, California; altitude 60 in ; precip- precipitation 100 mm. The site lies in the Colorado itation 100 mm. This saline aquatic site at the south end Desert near the town of Palm Desert. It consists of a of Death Valley represents aquatic ecosystems of the steep valley in which there is a creek that includes some Mohave Desert. It will permit comparisons with the sim- semipermanent pools. The vegetation is dominated by ilar environment of Locomotive Springs (Curlew Valley) Larrea divaricata and Opuntia spp. in the Great Basin. Rock Valley, Nevada; altitude 1,000 m; precipitation 100 mm. The site represents the Mohave Desert. It PROCESS STUDIES lies 80 miles northwest of Las Vegas and drains into the southern end of Death Valley. It is a broad bajada domi- It is not possible to determine a priori which eco- nated by Larrea divaricata and other shrubs. system constituents and which processes are likely to be Sycamore Creek, Arizona; altitude 600-1,000 m. most important in the dynamic functioning of the eco- This creek, which is a tributary of the Salt River, is an systems. Their relative importance will become apparent example of a watercourse in the southern deserts. Pre- only as the modeling activities proceed and after the cipitation in the lower part of the watercourse is 200- initial inventories for the validation study areas have 300 mm. been completed. The interactive nature of the program Tucson, Arizona; altitude 800-1,000 m. The implies that the list of processes and subsystems to be Sonoran Desert is represented by six areas ne.:r Tucson, studied must be constantly open to revision. However, a where rainfall averages about 250 mm. Four areas lie in tentative indication of the processes that are expected to the Santa Rita Experimental Range, where vegetation is require study are given below. In some cases it will be

28 30 possible to make use of data already in existence; for (other than predation), and reproduction. The possibility most of these processes, however, ad hoc studies will be of geographic differentiation within the species will be required. borne in mind. For each age and sex class, each process will be studied Terrestrial Ecosystems separately as a function of the various factors influencing it. MicrometeorologyFor each of the major community types occuri-ing in the sites of the validation studies, Soil Organisms For soil arthropods and perhaps nema- functions or submodels will be developed in which the todes, the questions to be answered by the process values of meteorological variables in the immediate vicin- studies do not differ from those for other animal groups. ity of the biota will be related to those measured in the For microorganisms, the approach may need to be mod- permanent recording stations. fled. It may not be practicable to perform regular cen- Special attention will be paid to distinguishable posi- suses of the populations, species by species. Instead it tions within the plant community where micrometeoro- may be necessary to treat the soil microflora collectively logical conditions are sufficiently different to constitute and consider its various types of activity (breakdown of a specialized niche. organic matter, nitrification and denitrification, nitrogen fixation, etc.) as functions of the controlling factors: Soil SubsystemsIn order that the conditions surround- temperatur.:, soil moisture, soil organic matter, aeration, ing plant roots and soil organisms may be adequately re- and composition of soil solution. The marked effects presented in the computer model, it will be necessary to that some microorganisms have on soil structure, partic- develop a submodel- by which the changes in physical and ularly on the crusting propertia of the soil surface, will chemical variables at particular points in the soil mass also be studied. can be calculated as functions of state variables in the ecosystem at large. A submodel is particularly desirable where meteorological inputs must be calculated. Aquatic Ecosystems Concurrently, detailed field observations will be made on the changes undergone by the physical and chemical A biotic Subsystems As with meteorological and soil parameters in certain soils. The observations will serve subsystems in terrestrial ecosystems, so studies on the as a check on the submodel and will identify improve- physical and chemical characteristics of aqueous environments that should be made. ments as functions of input and output will be needed. Additional process studies will cover the effects of Submodels will be required, for instance, of stream flow biota (plant roots, arthropods, burrowing vertebrates, patterns, of temperature distribution in flowing and microorganisms, lichens, and algae) on the physical and standing water, of changes in chemical composition dur- chemical properties of the soil. ing drying and refilling cycles, and the like. All these submodels will provide a means for relating the condi- PlantsFor each of the plant species important at one or tions within the ecosystem, which are immediately re- more validation studies, a series of processes will be levant to the functioning of the organisms in it, to the studied in order to design a model in which the abiotic imposed external conditions. environment can be represented. The possibilities of ecotypic or geographical variation in behavior within a Phytoplankton and Microorganisms As with soil micro- species will be borne in mind throughout. organisms, it may be necessary to treat the mixed popula- The processes studied will include water uptake and tion as a single collective entity, or in a few general transpiration, stomatal behavior, net photosynthesis, categories only, for the purpose of process studies. The growth and loss of aerial vegetative organs, root growth, functions to be studied will include photosynthesis, res- nutrient uptake, flowering and fruiting, and seed germ- spiration, nutrient uptake and biomass changes. ination and seedling establishment. The rate of each process will be expressed as a function of the variables Benthic and Littoral Plants The functions to be studied within the ecosystems that influence it. for these plants in the aquatic environment are in general the same as for terrestrial plants (see above), except that AnimalsFor each animal species OI group of species im- water relations are irrelevant. portant in one or more of the validation studies, the following processes will be studied insofar as appropriate: Animals The functions to be studied for aquatic animals Distributional pattern (including territorial behavior, -are the same as for terrestrial animals. Except for the quantity and composition of food consumed, change in fish, however, there is little knowledge of the relative biomass (including fat content), excretion, mortality importance of different animal groups in aquatic eco-

29 systems, and decisions on priorities for process studies results and will give the modeling group rapid access to will have to await the early results of modeling and valida- the data they need. tion studies. VALIDATION STUDIES MANAGEMENT Methodology D. W.Goodall is Director of the desert biome sub- Certain methodological requirements of the validation program and F. H. Wagner is Deputy Director. K. W. studies are being considered in ad hoc investigations. Bridges is Chief Modeler. Validation studies are grouped The direct measurement of the water balance of desert by sites, and each group is Li charge of a coordinator. ecosystems often poses difficulties. Accordingly, J. L. Similarly, process studies are grouped by subject- Thames is exploring an indirect approach based on sam- matter areas, and each group is in charge of a coordinator. pling of precipitation and soil moisture, D. F. Balph and The Director, the Deputy Director, the Chief Modeler, C. D. J orgensen are studying nondestructive methods of and the coordinators make up the Executive Committee. estimating populations of desert vertebrates, and G. L. The coordinators are identified in the Appendix (p. 31). Bender is responsible for similar studies of invertebrates.

Specific Areas RESEARCH METHODS AND PROGRESS Validation studies initiated during 1970 included the terrestrial studies at Tucson, a study of the temporary playa at Jornada, and the aquatic studies in Curlew Funding of this program began, on a restricted scale, Valley. in January 1970. A description of progress during the At Tucson, the four areas on the Santa Rita Experi- first six months of operation follows. mental Range have been fenced and treated; an elaborate meteorological station with remote continuous recording has been established; and initial inventories of plants, MODELING AND DATA PROCESSING vertebrates, and most invertebrate groups have been conducted. Some work on the biological inventories has Collaborators at each of the main study sites have also been performed on the Avra Valley bajada. (Project prepared word models of the systems with which they leader: J. L. Thames.) are concerned. These models describe the major struc- At Jornada, some preliminary records of vegetation tural and functional relationships between the compo- have been made on the whole catchment, but most ob- nents of the ecosystem, so far as they are known, and servations during 1970 have been limited to the playa. they will serve as a basis for developing mathematical The playa filled with water during heavy rains in July models for computer simulation. 1970, but prior to this time a full inventory of plants As a first step in the more formal modeling part of the and animals had been conducted. The rapid dcrelopment program, K. M. Bridges has developed a computer model of microorganisms, insects, and amphibia aftei the flood- for the Saratoga Springs ecosystem (see the list of valida- ing was followed in detail. (Project leader: W. G. tion studies, above). He used data collected by J. E. Whitford.) Deacon. In Curlew Valley, aquatic studies have begun in The major species occurring in the pond are repre- Locomotive Springs, at the south end of the valley, and sented as compartments in Dr. Bridge's model. Biological at four stations (spread over 30 km) in Deep Creek. At interactions between the species are considered as aspects each station, abiotic conditions are being monitored, and of energy flow between compartments. The model has a full inventory of algae, macrophytes, invertebrates, and been structured around an event chain that allows the fish has been conducted. (Project leaders: G. W. Minshall scheduling of discrete events. Approximate relationships and J. M. Neuhold.) for most of the energy-flow rates have been determined; hence it will be possible to check the model against the validation data. PROCESS STUDIES Facilities have been developed for the computer storage of all data collected in the biome operations, for pro- An important task during the first year has been that cessing the data, and for summarizing and retrieving of assembling data (published and unpublished) that may them. Thes,' facilities will enable collaborators to com- be relevant and deriving from early work in the validation municate rapidly with one another concerning research and modeling studies better guidance to the relative im-

30 9, portance of different processes in th:. -.;cosystem. How- of Nevada, Las Vegas (coordinator of validation studies, Saratoga Springs) ever, a few process studies were initiated in the field dur- G. R. Dutt, Department of Chemistry and Soils, University of ing the first year: Arizona, Tucson (Tucson validation studiessoil chemistry) Opuntia species in the Sonoran Desertphotosyn- J. H. Ehrenreich, Department of Watershed Management, Uni- versity of Arizona, Tucson (Tucson validation studies thesis, productivity, and water relations (D. T. Patten, vegetation) Arizona State University). E. Fish, Department of Watershed Management, University of Sitanion hystrix, an important perennial grass in Arizana, Tucson (Tucson -.alidation studiesplants) W. H. Fuller, Department of Agricultural Chemistry and Soils, the Great Basinphotosynthesis, root and shoot growth, University of Arizona, Tucson (Tucson validation studies and water relations (M. Hironaka and E. W. Tisdale, Uni- soils) versity of Idaho). R. W. Gordon, Department of Biology, Texas Tech University, Lubbock (Jornada validation studiesaquatic microorganisms) A triplex confertifoiia and Eurotia lanata, two Che- R. J. Hanks, Department of Soils and Meterology, Utah State nopodtaceous shrubs important in many parts of the University, Logan (coordinator of process studiesabiotic Great Basinphotosynthesis, root and shoot growth, and subsystems) D. M. Hendricks, Department of Agric,iltural Chemistry and water and nutrient relations (M. M. Caldwell and N. E. Soils, University of Arizona, Tucson (Tucson validation West, Utah State University). studiessoils) Rodents and lagomorphs in the Sonoran Desert M. Hironaka, Department of Range Management, University of Idaho, Moscow (ecophysiology of Great Basin perennial diets, fond preferences, and reproductive cycles (R. P. grasses) Balda, G. C. Bateman, and T. A. Vaughan, Northern A. J. Holman, Department of Wildlife Resources, Utah State Arizona University). University, Logan (Locomotive Springs validation studies aquatic invertebrates) Rodents of the Great Basinfood utilization and E. W. Huddleston, Department of En icmology, Texas Tech Uni- assimilation (D. R. Johnson, University of Idaho). versity, Lubbock (Jornada validation studiesaquatic insects) The role of algae in formation of soil crust (R. E. D. R. Johnson, Department of Biology, University of Idaho, Moscow (bioenergetics of small mammals) Cameron, California Institute of Technology). C. D. Jorgensen, Department of Zoology and Entomology, Biighani Young University, Provo, Utah (coordinator of validation studies, Pine Valley; methodology for estimates r)f small-mammal populations) APPENDIX: PARTICIPANTS R. H. Kramer, Utah Cooperative Fishery Unit, Utah State Uni- D. W. Goodall, Ecology Center, Utah State University, Logan versity, Logan (Locomotive Springs validation studiesfish) (Biome Director) M. L'Annunziata, Department of Agricultural Chemistry and D. Ashdown, Department of Entomology, Texas Tech University, Soils, University of Arizona, Tucson (Tucson validation Lubbock (Jornada validation studyaquatic insects) studiessoils) R. P. Balda, Department of Biology, Northern Arizona Univer- J. B. Low, Utah Cooperative Wildlife Research Unit, Utah State sity, Flagstaff (demography and food habits of small mam- University, Logan (Locomotive Springs validation studies mals) birds and mammals) D. F. Balph, Department of Wildlife Resources, Utah State Uni- Charles H. Lowe, Department of Zoology, University of Arizona, versity, Logan (coordinator of validation studies, Curlew Tucson (Tucson validation studiesreptiles) Valley; methodology for estimates of small mammal popu- J. A. Ludwig, Department of Biology, New Mexico State Uni- lations) versity, Las Cruces (Jornada validation studiesplants) S. A. Bamberg, Desert Research Institute, University of Nevada, R. I. Lynn, Department of Botany, Utah State University, Logan Reno (coordinator of process studiesplants) (Locomotive Springs validation studiesphytoplankton) G. C. Bateman, Department of Biology, Northern Arizona Uni- A. N. MacGregor, Department of Agricultural Chemistry and versity, Flagstaff (demography and food habits of small Soils, University of Arizona, Tucson (Tucson validation mammals) studiessoil microorganisms) G. L. Bender, Department of Zoology, Arizona State University, Clark Martin, Watershed Management, Cooperative Extension Tempe (methodology for estimates of invertebrate popula- Service, University of Arizona, Tucson (Tucson validation tions) studiesSanta Rita) C. D. Bonham, Department of Watershed Management, Univer- G. F. Meenaghan, Department of Chemical Engineering, Texas sity of Arizona, Tucson (Tucson validation studiesplants) Tech University, Lubbock (Jornada validation studiesabiotic K. W. Bridges, Wildlife Resources, Utah State University, Logan conditions in playas) (Chief Modeler) G. W. Minshall, Department of Biology, Idaho State University, D. R. Cable, Watershed Management, Cooperative Extension Pocatello (coordinator, Aquatic Process Studies; coordinator, Service, University of Arizona, Tucson (Tucson validation Deep Creek validation studies) studiesSanta Rita) J. N. Minshall, Department of Biology, Idaho State University, M. M. Caldwell, Department of Range Science, Utah State Uni- Pocatello (Deep Creek validation studies) versity, Logan (photosynthesis of Great Basin shrubs) J. M. Neuhold, Ecology Center, Utah State University, Logan R. E. Cameron, Jet Propulsion Laboratory, California Institute (Locomotive Springs validation studies) of Technology, Pasadena (algal crusts on soils) W. L. Nutting, Department of Entomology, UMversity of Robert M. Chew, Department of Biology, University of Southern Arizona, Tucson (Tucson validation studiesinsects) California, Los Angeles (coordinator of process studies D. T. Patten, Department of Botany, Arizona State Univeisity, vertebrates) Tempe (ecophysiology of cacti) E. L. Cockrum, Department of Biological Sciences, University F. J. Post, Department of Bacteriology, Utah State University, of Arizona, Tucson (Tucson validation studiesmammals) Logan (Locomotive Springs validation studiesmicrobiology) G. L. Cunningham, Department of Biology, New Mexico State H. K. Oashu, Department of Hydrology and Water Resources, University, Las Cruces (Jornada validation studiesplants) University of Arizona, Tucson (Tucson validation studies James E. Deacon, Department of Biological Sciences, University hydrology)

31 Ralph .1. Raitt, Department of Biology, New Mexico State Uni- versity, Las Cruces (Jornada validation so liesbirds) H. G. Reynolds, Rocky Mountain Forest and Range Experiment Station, U.S. Department of Agriculture, Arizona State University, Tempe (Tucson validation studiesmammals) S. M. Russell, Curator of Birds, University of Arizona, Tucson (Tucson validation studiesbirds) II. D. Smith, Department of Zoology, Brigham Young University, Provo, Utah (methodology for estimates of small-mammal populations) E. E. Staffeldt, Department of Biology, New Mexico State Uni- versity, Las Cruces (coordinator of process studiesmicro- organisms) 180' C. B. Stalnaker, Utah Cooperative Fishery Unit, Utah State Uni- versity, Logan (Locomotive Springs validation studiesfish) J. L. Thames, Department of Watershed Management, University of Arizona, Tucson (coordinator of validation studies, Tucson) Irvin P. Ting, Department of Life Sciences, University of Cali- fornia, Riverside (coordinator of validation studies, Deep Canyon) E. W. Tisdale, Department of Range Management, University of Idaho, Moscow (ecophysiology of Great Basin perennial grasses) F. B. Turner, Laboratory of Nuclear Medicine and Radiation Biology, University of Calif-Nrnia, Los Angeles (coordinator of validation studies, Rock Valley) T. A. Vaughan, Department of Zoology, Northern Arizona Uni- FIGURE 7Circumpolar areas designated as sites for in- versity, Flagstaff (demography and food habits of small tensive tundra research. The areas and the nations having mammals) jurisdiction over them are as follows: Devon Island (Cape F. II. Wagner, Department of Wildlife Resources, Utah State Sparbo) and Mackenzie Delta, Canada; Kevo, Finland; University, Logan (Deputy Biome Director) Moor House, Great Britain; Disko Island, Greenland C. R. Ward, Department of Park Administration, Horticulture (Denmark); Glenamoy, Ireland; Hardangervidda, Norway; and Entomology, Texas Tech University, Lubbock (Jornada Abisco, Sweden; Barrow, Alaska, United States; Niwot validation studiesaquatic insects) F. G. Werner, Department of Entomology, University of Arizona, Ridge, Colorado (not shown on map), United States; Tucson (coordinator of process studiesinvertebrates; Tucson Taimyr Peninsula (Pyasina River) and Salekhard (Polar validation studiesinsects) Urals), Union of Soviet Socialist Republics. N. E. West, Department of Range Science, Utah State University, Logan (ecophysiology of Great Basin shrubs) W. G. Whitford, Department of Biology, New Mexico State Uni- cally to seemingly minor perturbations. This response is, versity, Las Cruces (coordinator of validation studies, Jornada) in part, the result of increased energy flow into the upper W. H. Woodin, Arizona-Sonora Desert Museum, Tucson, Arizona substrate as the insulating covers of vegetation and peat (Tucson validation studiesreptiles) are grazed, physically disrupted, or removed. The phys- James R. Zimmerman, Department of Biology, New Mexico State University, Las Cruces (Jornada validation studies ical consequences are acceleration of thaw and rapid insects) erosion of the underlying ice-rich permafrost. These consequences are easily observed and measured. However, the biological responses to disturbances and to steps taken to reduce them are not easily observed, and their consequences are not immediately known. Thus, it is TUNDRA BIOME essential to investigate these responses. Initial changes must be studied, and cumulative effects must be anticipated and measured. The main purpose of the U.S. tundra biome program Many biological responses to disturbances in the arctic is to analyze the structure and function of the wet arctic and subartic ecosystems are related to warmer surface tundra. The research is designed to develop a predictive and substrate conditions and increased availability of understanding of the wet tundra ecosystem and will em- nutrients. Increases in primary production during the phasize the application of environmental knowledge to short growing season result from rapid initial release of the problems of degradation, maintenance, and restora- nutrients in the physically disturbed organic layers and tion of these ecosystems. from release of previously inaccessible nutrients in the Numerous international sites, most of which are in the lower root zones and thawing permafrost. Initial modifi- circumpolar region, have been selected (Figure '7). An al- cations in the nutrient and energy flows can be followed pine site in the Colorado Rockies is included in the U.S. by comparing, at both disturbed and undisturbed sites, program. such parameters as available nutrients, metabolic activi- The arctic and subarctic ecosystems :espond dramati- ties, population dynamics (microorganisms and larger

32 4: consumers), and thermal regimes in the vegetative and ABIOTIC VARIABLES soil covers. INCOMING ENERGY

mmas mg Moi u e

RIDGES LOWLAND MEADOWS THAW and POLYGONAL...and LAKES RESEARCH PLANS SLOPES GROUND MARSHES

I PONDS 1 if

During the summer of 1970, an integrated bioenviron- NUTRIENT POOLS mental program was begun in northern and central Alaska. Barrow, the intensive site, and several compara- BIOTIC VARIABLES tive sites are situated along a bioclimatic gradient rang- PRIMARY PRODUCERS ing from the Arctic Ocean to the taiga of interior Alaska.

This research is designed to evaluate temporal and spatial TERRESTRIAL AQUATIC HERBIVORES variations in rates and patteins of nutrient cycling and DECOMPOSE RS DECOMPOSERS energy flow in natural and perturbed arctic and subarctic environments. Also evaluated are the effects of man- made disturbances and simulated natural stresses on nu- CARNIVORES trient cycling and energy flow through the major components of cold-dominated arctic and subarctic ecosystems. FIGURE 8General model of wet tundra ecosystem. The research effort during 1970 emphasized comparisons between natural and disturbed ecosystems, comparisons that were designed to establish an earl:y understand- Modeling activities are an integral part of the research. ing of how sensitive the ecosystems are to stresses and of A primary production model for the tundra canopy is how they adjust after being stressed. Two types of exper- being developed at San Diego State College. A gross air- iments involving disturbances in ecosystems have been land meterological model is being developed by the established at the Barrow, Alaska, site: artificial and nat- Center for Environment and Man of the University of ural. The artificial disturbances consist of simulation of Connecticut. The U. S. Army's Cold Regions Research anticipated man-made disturbances on the temperature- and Engineering Laboratory is perfecting a thaw model sensitive permafrost terrain. These include (1) increased that will be used to preciict soil temperature and depth of soil temperatures to simulate heat flow out of the tundra thaw. A general tundra ecosystem model was initiated in over a buried hot oil pipeline, (2) increased air tempera- the fall of 1970. ture and reduction of wind speeds in the plant canopy to Central services for the program are provided through test the response of fauna and flora to more favorable the University of Alaska. These include chemical analyses gross,ing conditions, (3) physical disturbance of the sur- for plant and soil samples and data processing. The data face vegetation, and (4) oil spills of different intensities acquired :n previous years from Barrow are being pro- and duration on both land and fresh water. Validation of cessed at San Diego State College. Logistic support in a precision soil-thaw model is being conducted on each northern Alaska is provided by the Naval Arctic Research treatment and on control sites. Laboratory, Barrow, Alaska. The natural experiments simulate naturally occurring J. Brown is Biome Director and G. C. West is Deputy phenomena that affect the rate of nutrient and energy Director. The organizational structure is shown in the flow through the ecosystem (i. e., soil-plant-air processes). following list: These include (1) clipping of vegetation to simulate increased animal grazing, (2) mulching of natural plant A. Central program litter to simulate accumulatinn of organic matter, and (3) 1. Executive fertilization of both terresti:J and aquatic systems. a. Director A variety of physiological processes are being measured b. Deputy Director on sites for experiments of both types. The processes in- 2. Services clude plant and animal productivity, microbial activity, a. Administrative life cycles of insects, and soil and air temperature b. Data processing regimes. c. Data analysis Each disturbed site is being compared with an undis- d. Data banks turbed control site. The design is organized according to e. Analysis and modeling the four basic ecosystem components illustrated in f. Sample analysis Figure 8. g. Editorial

33 3. Scientific coordination Ponds, an integral aspect of ail principal habitats, show a.Producers various stages of succession resulting from both filling b.Consumers and erosion. Large shallow lakes occupy 30% of the area north of 710 latitude. These vegetation, soils, microrelief c.Microbiology and aquatic environments form a complex mosaic which d.Nutrient flux influences plant productivity and animal population and e.Abiotic diversity throughout the year. Ridges, polygon tops, and f.Aquatic slopes commonly contain in decreasing order of occur- Biome-wide rence Carey aquatilis, Petasites frigidus, Salix pulchra, g. Arctagrastis latifolia, Poa arctica, Luzula con fusa, Salix h. Analysis and modeling rotundifolia, and Eriophoruin scheuchzeri. Dupontia i.National, international fisheri, Carex aquatilis, Eriophorum anqustifolium and B. Research program E. scheuchzeri, Petasites frigidus, and Poa arctica occur 1. Intensive site studies in meadows and wet polygonal troughs. Arctophilo fulva a. Producers is found in shallow ponds. Primary production is controlled by a combination of b. Consumers temperature, soil moisture, day length and nutrient c. Decomposers supply. The more productive habitats on micro- and d. Abiotic meso-scales are near the wetter end of the soil moisture e. Aquatic gradient (meadows and polygon troughs). All habitats 2. Biome-wide studies are controlled by the presence of permafrost (0° C a. Arctic and alpine, Alaska ground temperature or colder) near the surface throughout the growing season. Especially significant to the (1)Producers amount of seasonal primary production are the climatic (2)Consumers conditions of the first two or three weeks of the growing (3) Decomposers season and the previous summer's stored reserves and (4) Abiotic climate. Superimposed on this are fluctuations in nu- b. Alpine, Colorado trient cycle. On all terrestrial sites, but particularly on the wetter meso-sites, production is directly influenced (1) Producers by delayed effects of decay of clipped plant debris and (2)Consumers fecal remains. It is hypothesized that the availability of (3) Decomposers nutrients to plants for growth is modulated by the lem- (4) Abiotic ming population through accumulation and storage of certain elements (N, K, P) in, and their ultimate release from, fecal and other dead oronic matter. Long-term rates of total decomposition in most of RESEARCH METHODS AND PROGRESS the habitats are approximately equal to production; there is no net accumulation of organic matter. Initial organic matter accumulation on drained lake beds is rapid. Based Field studies in Alaska were initiated in June 1970. on evidence, depth of thaw measurements, and the ab- About 100 scientists and technicians were involved. sence of widespread organic terrain, organic accumulation and decomposition under uniform climatic conditions In order to analyze the great amount of data emanat- are assumed to reach a steady state within a relatively ting from these studies, a central data-processing system short period (presumably less than 100 years). A domi- has been established at the University of Alaska. Central- nant acceleration effect in the decomposition process is ized laboratory analysis and data processing provide the amount of standing dead plant matter prostrated by lemming grazing. rapid turnaround of statistical reports, samples, and data. Photosynthesis on a daily basis is efficient due to the Data storage facilities have been developed for research long photoperiod and generally low temperatures. During projects that were active at Barrow, Alaska, over the last the "night," however, and particularly in the lower levels 25 years. A summary data report was published in July of the canopy, carbon dioxide uptake is limited by the 1970. The Barrow data have been synthesized into a availability of light. Grasses all show a high carbon dioxide compensation concentration, a pronounced oxygen word model, or description, of the Barrow site. The fol- inhibition, and low light requirement for saturation. lowing model, constructed by project participants, is Within species, photosynthesis serves as a function of being continuously refined: leaf position, age, and season. Prominent saprovores are dipterous larvae, collembo- The arctic tundra near Barrow, Alaska, encompasses a lans and oribatid mites. Although the rate of organic de- complex of habitats arrayed along a moisture-dominated composition is slow, wetter habitats show higher rates of gradient. Vegetation and soils are distributed along nutrient loss through decomposition compared to better- micro-, meso-, and macro-environments according to type drained, drier habitats. Microbial activity appears to be and size of polygons, regional relief, and land form type. several orders of magnitude below that of lower latitude These range from upland meadow communities (arctic wet ecosystems and, furthermore, is confined to the brown and upland tundra soils) to wet meadow and upper 10 centimeters or so of the soil. marsh types (meadow and bog soils) to emergent aquatics There are low numbers of thermophilic bacteria in (hydrosols) and open water in small ponds and lakes. most arctic soils. Mesophilic micro-organisms are greatest

34 in numbers when the soil is frozen and drop sharply once Productivity and Conservation in Northern Circumpolar the summer thaw occurs. Psychrophilic bacteria hcrease Lands, Edmonton, Alberta. Canada, Oct. 15-17,1969. in numbers during the summer. Limited metabolic activ- International Union for Conservation of Nature and Mtural ity also occurs during the summer. Resources, Morges, Switzerland. Coulombe, H. N., and J. Brown. 1970. Analyses of the structure important for the assessment of production on tundra and function of the wet tundra ecosystem at Barrow, Alaska. is the distinction between aboveground and underground Abiotic data report, 1965. Bureau of Ecology, San Dicgo components of plants. Biomass ratios below to above are State College, San Diego, Calif. of the order 5:1 or greater. Annual production rates Brown, J., and G. C. West. 1970. Tundra biome research in below to above are approximately equal. Alaska: the structure and function of cold-dominated eco- Annual production rates for the tundra ponds and the systems. Tundra Biome Report 70-1. U.S./International lakes studied are very low compared to most temperate- Biological Program. zone water. Many of these small ponds receive seasonal influxes of organic matter (clippings, faeces, soluble organic), especially during the spring runoff. During the APPENDIX B: PARTICIPANTS summer, thermal erosion of organic-rich permafrost also contributes to lake and pond filling. Approximately 25% J. Brown, Cold Regions Research and Engineering Laboratory, U.S. Army, Hanover, N.H. (Biome Director; thaw validation; of the total seasonal lake production occurs beneath the ecosystem analysis) ice before and after the spring thaw and after freeze-up. V. D. B. Alexander, Institute of Marine Science, University of Production appears limited by a short growing season, Alaska, College, Alaska (phytoplankton; nitrogen fixation) low solar intensity during the growing season and nutri- J. H. Anderson, Institute of Arctic Biology, University of ent deficiencies, especially for phosphorus. Alaska, College, Alaska (producers) Plant consumption is dominated by the brown lem- R. J. Arkley, Department of Soils and Plant Nutrition, Univer- ming (Lemmus trimucronatus), which utilizes standing sity of California, Berkeley (pedology and nutrient cycling) living tissue on a year-round basis. Other vertebrate her- R. J. Barsdate, Institute of Marine Sciences, University of bivores typical of tundra elsewhere are here negligible Alaska, College, Alaska (phytoplankton; nutrient cycling) R. E. Benoit, Department of Biology, Virginia Polytechnic Insti- (e. g. a second microtine, Dicrostonyx; caribou; geese) or tute, Blacksburg (microbial activity) absent (ground squirrels). The herbivory of arthropods is C. S. Benson, Geophysical Institute, University of Alaska, Col- probably lower than in other ecosystems. The vegetation- lege, Alaska (micrometeorology) lemming interaction is cyclic on a "short-term" basis, H. N. Coulombe, Biology Department, San Diego State College, the period being three to five years. San Diego, Calif. (ecosystem analysis) One type of carnivory is conspicuously associated with P. W. Flanagan, Botany Department, University of Alaska, Col- the lemming cycle, with several bird and mammal pred- lege, Alaska (microbial activity) ators present in significant densities. These include jae- P. L. Gersper, Department of Soils and Plant Nutrition, Uni- versity of California, Berkeley (pedology; nutrient cycling) gers, owls, and gulls (summer active), and weasels and N. W. Lord, The Center for Environment and Man, Inc., Hart- foxes (active year-round). Other carnivory is that of pred- ford, Conn. (air-land interface modeling) ators on insects, mainly birds but also including shrews, S. F. MacLean, Jr., Department of Zoology, University of Illinois, arachnids, and other arthropods. Most conspicuous is the Urbana (soil arthropods) carnivory of shorebirds which is concentrated on Diptera B. McCoun, Cold Regions Research and Engineering Laboratory, larvae. U.S. Army, Hanover, N.H. (plant physiology; oil ecology) Dominating all terrestrial and aquatic processes is the P. C. Miller, Biology Department, San Diego State College, San shortness of the growing season, varying between 45 and Diego, Calif. (primary production model) 90 days. Short growing season is partially compensated Y. Nakano, Cold Regions Research and Engineering Laboratory, U.S. Army, Hanover, N.H. (thaw validation) for by uninterrupted diurnal light. J. P. Pandolfo, The Center for Environment and Man, Inc., Hartford, Conn. (air-land interface modeling) Perturbation studies have stressed the system in such F. A. Pitelka, Department of Zoology, University of California, a way that physical and biological responses are evident. Berkeley (herbivore populations) A. Schultz, Department of Forestry, University of California, Preliminary analyses indicate different species sensitivi- Berkeley (pedology; nutrient cycling) ties to oil spills, a marked growth response as a result of L. L. Tieszen, Biology Department, Augustana College, Sioux increased air temperatures, and increased production as Falls, S. Dak. (photosynthesis; primary production) K. Van Cleve, Department of Forestry, University of Alaska, a result of fertilization, especially with phosphorus. Re- College, Alaska (nutrient cycling; pedology) sponses associated with older physically disturbed areas G. E. Weller, Geophysics Department, University of Alaska, appear to be increased rates of nutrient and energy flow, College, Alaska (micrometerology) G. C. West, Institute of Arctic Biology, University of Alaska, since production is frequently seen to be high while College, Alaska (Deputy Biome Director; herbivore popula- amounts of standing dead and litter are low. All these tions) are accompanied by an apparently enhanced activity of microbial decomposition.

TROPICAL FOREST BIOME APPENDIX A: PUBLICATIONS Brown, J., F. A. Pitelka, and H. N. Coulombe. 1970. Structure and function of the tundra ecosystem at Barrow, Alaska: a Biological productivity levels in tropical forests are word model, p. 41-71. In Proceedings of Conference on among the highest known, although soil nutrient levels

35 are extremely low. Studies of tropical forests are impor- The Biome Director is H. T. Odum, Department of tant for two main reasons: Environmental Engineering, University of Florida, These forests are being rapidly destroyed by log- Gainesville. Members of the steering committee are as ging and by slash-and-burn agriculture. The destruction follows: often results in a near-desert, be:..ause the soil nutrients are removed with the trees. Richard S. Cowan, Smithsonian Institution, Washington, Knowledge of tropical forests is essential to under- D.C. standing the ecosystems of the world. The forests repre- R. S. Davidson, Battelle Memorial Institute Columbus sent extremes for many environmental and biological Laboratories, Columbus, Ohio parameters. Troy C. Dorris, Reservoir Research Center, Oklahoma The research design will be similar to those outlined State University, Stillwater for the other biome studies in the analysis-of-ecosystems Frank B. Golley, Institute of Ecology, University of program. Georgia, Athens Several organizational and planning meetings were Thomas E. Moore, Museum of Zoology, University of held during 1969. Site surveys have been made in Central Michigan, Ann Arbor and South America. A variety of sites are available in Jack T. Spencer, The Organization for Tropical Studies, Guatemala, Costa Rica, Panama, Colombia, Venezuela, Inc., North American Office, South Miami, Fla. and Puerto Rico. U.S. groups interested in cooperation F. H. Wadsworth, Institute of Tropical Forestry, Forest with the 1BP are working with scientists from these Service, U. S. Department of Agriculture, Rio Piedras, countries. Puerto Rico

36 Biological Productivity in Upwelling Ecosystems

The major features of an oceanic ecosystem are deter- Oceanographic cruisesto Peru in 1969 and to the mined for many regions by the vertical mixing rate, that Mediterranean 4,1 1970 is, by the rate at which nutrient-depleted water in the Projectsone on construction of a simulation eutophic zone is replaced by nutrient-rich water from model and one on applicaticns to aquiculture below. In tropical and subtropical regions these rates are Conferences and symposia often low; as a result, there is little biological produc- Information obtained from the cruises is vital to the tivity and blue water. However, along the west coasts of functioning of the projects on which activity is expected the continents the surface water is driven offshore by to focus in the later phases of the program. the trade winds and is replaced by deep water. The term Significant contributions already have been made to "coastal upwelling" is applied to this condition. this program by the cooperating Mediterranean labora- Regions affected by coastal upwelling are highly pro- tories. The arduous task of obtaining and analyzing the ductive, often supporting large fisheries. An understand- phytoplankton data has been undertaken by personnel ing of the dynamics of production in these regions would of the Instituto de Investigaciones de Pesqueras in Bar- contribute to many marine activities, including manage- celona. A program on ISN productivity measurements in ment of fisheries and aquiculture, pollution control, and the Mediterranean by the Station Marine d'Endoume in artificial upwelling. Marseile is continuing as a result of participation in the The Program Director is Richard C. Dugdale. Peruvian cruise. Several Greek scientists are expected to participate in the Mediterranean cruise. The Greek scientists are from the Greek Hydrographic Service, the Dem- ocritos Nuclear Research Center, and the Department of OBJECTIVES Botany of the University of Athens. Opportunities for exchange of personnel between laboratories are inherent in the upwelling program. For Objectives are: example: To obtain sufficient quantitative understanding Facilities are being made available at the Instituto of the structure and dynamics of upwelling ecosystems de Investigaciones Pesqueras in Barcelona. to allow prediction of the consequences of perturbations A visiting lecturer program is in effect at the Sta- of these and other marine ecosystems when these changes tion Marine d'Endoume in Marseille. result primarily in enhanced circulation of nutrients into The Progam Director has been invited to lecture lighted regions of the sea in Greece on elementary simulation modeling techniques. To apply the theoretical concepts so obtained to A graduate student from the United States will pilot aquicultural projects work several months at the University of Athens, where To construct a simulation model of the Peruvian a program on plant protein production is under way. anchovy fishery, incorporating the dynamics of the nutrient concentration and primary productivity CRUISES

The program has been designed to be truly interna- RESEARCH PLANS tional and yet limited in scope and geography so that important basic information can be obtained in a relatively short time with a modest expenditure of funds. The program encompasses scientific activities in three Two contrasting areas have been selected: the Peruvian categories: coastal upwelling system and the Mediterranean Sea. The

39 importance of the Peruvian area arises from the intense wherein populations of microorganisms are held in a re- upwelling and the associated anchovy fishery. The Med- actor under nutrient limitation that controls the rate of iterranean, in contrast, is impoverished in nutrients but biological production. In a true chemostat, nutrient supports a fishery of importance to the surrounding medium is fed into the reactor at a constant rate, and countries. Little is known of the circulation of nutrients nnediurn from the reactor containing the organisms is ex- in the Mediterranean, but local areas of enhanced vertical pelled at the same rate. In a chemostat at steady state, circulation occur that are vital to the maintenance of the growth rate of the organisms is set by the feed rate biological productivity. and the concentration of the organisms is set by the con- The technique of applying identical methods to concentration of limiting nutrient in the feed. The produc- trasting environments, a highly effective means of study- tivity is regulated by the amount of incoming limiting ing ecological systems, will be exploited fully. nutrient. Scientists from the United States, Peru, France, The chemostat technique shows great potential for Greece, and Spain are participating in the program. use in some phases of aquiculture, because control of production rate and species composition is possible. However, chemostats normally run under closely regulated operat- SIMULATION MODEL PROJECT ing conditions, including temperature, contamination, and light in the case of algae. The similarity between The simulation model project is based at the Univer- chemostat conditions and those of phytoplankton pop- sity of Washington, Seattle, where courses in simulation ulations in the tropical and subtropical oceans suggests techniques are taught at the Center for Quantitative that it might be possible to approach the problem of Science and in the Department of Oceanography. The algal production in pon-ls and lagoons with chemostat project supports travel and maintenance for investigators procedures. Since significant departures from the rigorous who wish to go to the University of Washington to learn conditions of true chemostat operation would occur, simulation and computer techniques or to work on a these experimental enclosures probably should be con- model. sidered quasichemostats. Since 1963, several commercial fisheries have used The first successful attempt to operate algal quasichem- large-scale digital computer simulation models to evaluate ostats was made by two students in a phytoplankton biological and economic consequences of varic is manage- ecology course at Friday Harbor Laboratories in 1968. ment policies, but there has been no previous attempt to They used plexiglas reactors with a volume of roughly construct a simulation model for the Peruvian anchovy 100 liters and water from the laboratory seawater sys- fishery. Detailed analytical models have been used to es- tem. Flow rates, nutrient concentration in the seawater timate equilibrium yield and to study the recruitment feed, and light conditions varied considerably. They were processes of the anchovy. The biological statistics col- able to attain a quasi-steady-state in which nutrient con- lected for these analytical models provide a data base for trol by silicate was clearly evident. The flow rates were constructing a comprehensive simulation model of the set to give doubling times of about 24 hours in one re- anchovy stocks. actor and about 36 hours in the other; both were initially Past attempts by oceanographers and fisheries scien- seeded from the contents of a net tow from surface water. tists to construct models of upwelling ecosystems have Most added species washed out at these high flow rates, been artificial and oversimplified. Many factors, particu- and the ultimate populations were about as follows: larly the lack of well-coordinated fisheries and ocean- Skeletoneina costatum, 90 percent; and 6 species of ographic studies of marine ecosystems, have obstructed Chaetocerus, 10 percent. progress in this area. Recent developments i1 simulation These results warrant further investigation into the modeling on digital computers present new opportu- characteristics of "open-sky quasichemostats." It is plan- nities for constructing quantitative models of marine ned to repeat the Friday Harbor experiments during the ecosystems. anchor station on the Second Thompson cruise off the coast of Peru. The extent to which the chemostat system would AQUICULTURE PROJECT have immediate value in aquiculture would depend on its adaptability to large containers, tanks, and eventually Efficient aquiculture requires prediction and control pools, and on its independence of artificial radiation and of quality and quantity of the biological material being temperature control. Long-range aquicultural uses include produced. An important step is to control the rate of pri- the application of chemostat principles in the utilization mary production and the species composition of the of artificially upwelled deep water and use of treated algae in photosynthetic systems. sewage or other pollution products in natural or man- A neglected approach is to use chemostat techniques made enclosures or impoundments.

38 40 CONFERENCES AND SYMPOSIA the direction was consistently from the southeast, that is, parallel to the coast and directly opposite to the flow The conferences are required to maintain direction of water at the 25-m depth. and momentum during the entire period. The more These findings, together with the results of a number formal symposia are to be held later for the dissemina- of surface drogue experiments, give a consistent picture tion, discussion, and synthesis of results that are well of an advancing and retreating front in the region studied. beyond the preliminary stage. The front advances to the north as the winds blow more strongly and retreats to the south as the wind stress is relaxed. Surface flowing water is accelerated to the left RESEARCH METHODS AND PROGRESS and tends to move offshore to be replaced by southerly flowing water from about 25 m, which is simultaneously accelerated in the onshore direction. It is likely that bot- CRUISE tom topography has some effect on this simplified model of water movement. The persistently low temperatures In early 1969 a cruise was made to the coast of south- along lines 6 and 8 (Figure 1) during most of the study ern Peru. A rectangular grid 15 miles long and 30 miles are striking and may be caused by a submarine ridge that wide was established about 5 miles off the Peruvian coast extends out from the shoreline. (see Figure 1). The grid area was studied intensively during two visits of the Thompson and on three shorter Nutrient Circulation visits of the Gosnold. Standard productivity stations were made throughout During the first visit, an intensive survey was made of the survey and included the measurement of primary the grid area and three strings of current meters were set. productivity by 14C uptake, chlorophyll and light mea- Routine data were processed by the shipboard data reduc- surements, phytoplankton sampling, and the measure- tion system that includes a computer and an on-line data ment of nutrient demand through measurements of15N- acquisition system. Automatic sampling instruments labeled nitrate and sometimes ammonia. The measure- measured the temperature, salinity, and the concentra- ment of nitrate demand by the phytoplankton population of chlorophyll, nitrate, and silicate of water pumped tion a.t all the survey stations makes it possible to com- from three meters depth while the ship passed over a pute ar, average upwelling velocity if the nitrate content predetermined path on the survey grid. Also incident of the upwelling source water is known. A preliminary radiation, wind speed and direction, and ships heading estimate of the upwelling rate by this means is about 0.5 and direction were measured and recorded. From these to 1 m per day. data maps were produced that showed the distribution During Leg 2, an effort was made to determine the of temperature or chlorophyll or the two nutrients. kinetics of phytoplankton nutrient uptake. These studies About 24 hours of sampling time were required to pro- required the cooperation of several scientists. Five scien- duce maps of the entire region. Maps of smaller areas tists studied nitrogen uptake and three studied silicate valuable were made during the night. These maps were uptake. during the second visit because they were used in plan- The study of nitrogen uptake included the study of ning the acquisition of water samples for experimental nitrite, nitrate, and ammonia. Reaction rates of nitrate and purposes. ammonia uptake were obtained as functions of light in- Groups of scientists collaborated on various problems. tensity and nitrate and ammonia concentration. The ef- Information was exchanged during evening seminars. fect of light intensity on nitrate and ammonia uptake Before the end of the cruise most of the data were re- can be described by the Michaelis-Menton expression duced to a form suitable for initial analysis. with great accuracy. The constants for the Michaelis- Menton expression do not vary significantly from the Physical Oceanography 100 percent light level to the 10 percent light level. At All the current meters and the wind instrument carriedhigh nitrate concentrations, nitrate uptake was found to on the two strings recovered at the end of firstthe visit be proportional to nitrate concentration. Measurements were found to have functioned while in place. All records made in a shipboard chemostat have been used to derive have been read and processed. The results show that the a quantitative expression for the suppressiveeffect of currents at depths greater than 25 m were to the south- ammonia on nitrate and nitrite uptake. east, that is, parallel to the coast. Once during the cruise Nitrate uptake was also studied in relation to nitrate this southerly flow ceased, and a temporary, small north- reductase activity. Two scientists collaborated in stand- erly component developed. The wind data showed clearly ardizing their procedures for measuring the activity of that although there were large fluctuations in velocity, nitrate reductase. Their subsequent studies revealed a

39 40' 76° 40' o' 75° MNI.Elm IMMON411M MI1=-Mi

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8 -6 = 3,7,10,32,48 II

34 53 H 24 8 88,35,37

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Pta. San Juan 21 0.#3.4642 0 tit0 4$1

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18 16° FIGURE 1Cruise track for R/V Thompson 36. The letters A, B, C, and D designate lines of grid parallel to the coast; CM designates current meter locations; circled numbers designate lines normal to the coast; other numbers designate hydrostations. strong daily cycle in nitrate reductase activity in the depletion of silicate in carboys exposed to sunlight. They phytoplankton population. A comparison of the nitrate also made partiwiate silicate measurements. The deter- reductase activity with nitrate uptake suggests the a mination of isotope ratios in the particulate silicate frac- second nitrate reductase is present that is not being tion is laborious and time-consuming, and the results are measured. not yet available. If the water is relatively stable and the A coalition of scientists studied silicate cycling. They phytoplankton population has been present some days, measured the uptake of silicate by phytoplankton by then the particulate matter has a nitrate-silicate ratio using the heavy isotope 28Si and by ob.Aving the rate of that closely matches the nitrate-silicate ratio in the water.

40 42 This similarity suggests that silicate, like nitrate, was on shipboard by one of his students. A considerable num- taken up by diatom populations in amounts that were in ber of samples were counted on shipboard, and a running proportion to the amounts available in the water. In the account of gross species composition was made. How- case of nitrate, it appears that nitrate reductase, an in- ever, most samples have been returned to the Instituto ducible enzymP, is synthesized in proportion to the de Investigaciones Pesqueras in Barcelona for counting amount present in the water, providing an explanation and analysis. During a visit with the Program Director for the observed rnult. Finding an explanation for sili- after the cruise, plans were laid for analyzing the data cate uptake may be more difficult. Nevertheless, the in- and for next year's work. The productivity and hydro- formation on nitrate and silicate uptake has important graphic data will be used to perform multiple correlation implications for control of the nutrient ratios in the up- analysis on a computer. welling area. An important consideration in the analysis of the spe- Maps of the nitrate-silicate ratio at 3 rn in the study cies composition may be the instantaneous growth rates area show large differences. The differences appear to of the population. These rates have already been com- arise from (1) the depth of the upwelling water, a result puted from the data on nitrate and ammonia uptake, and of a depletion in silica relative to nitrate in the region another valuable contribution may be made when RNA- above the pycnocline, and (2) the appearance of dino- DNA analyses are completed. flagellate blooms that leave silicate untouched while consuming nitrate. This ratio may be useful in assessing the Food Chains depths from which the upwelling water is derived am. ; a means of distinguishing between diatom and dinoflagel- Preliminary processing of the zooplankton collections late activity. has been completed at Woods Hole Oceanographic Insti- Although phytoplankton populations in recently up- tution. Settling volumes have been measured and the col- welled water are often believed to show a lag period be- lections have been sorted into major groups. Further fore normal growth takes place, the phenomenon was studies of these groups will be made. A program of labo- not observed on the Anton Bruun 15 or during the major ratory culture of species of Pseudocalanus has been insti- part of the Peruvian cruise. Work carried out on the tuted. The zooplanktors have been carried through an Thompson has been very helpful in explaining this ap- entire generation. They appear to require freshly pro- parent conflict. Factorial experiments were conducted in duced phytoplankton, without which the individuals die which various chelating agents were added to samples of or do not produce eggs. The females produce egg snacks seawater. The effects wcre assayed with 14C. When lo- as rapidly as one every 2 days, and one female may pro- calized upwelling prevailed, the recently upwelled water duce as many as 200 new individuals in 2 weeks. was chelated by naturally occurring organic compounds The research on excretion products as a means of to such an extent that no increase in growth rates was in- distinguishing and quantifying grazing activity by zoo- duced in the experiments. However, when massive up- plankton and by fish is well under way. A reliable welling flooded the study area, the water was poorly che- method for the analysis of creatine in seawater was de- lated and higher growth rates were experimentally veloped and used on the Peruvian cruise. Ordinarily, cre- induced. The working hypothesis is that sufficient iron atine could not be detected in seawater except under is always available in surface seawater but that its avail- conditions of high fish concentration. In several experi- ability depends on the presence of naturally occurring ments, anchovies were caught and placed in holding chelating compounds. Presumably, the chelators are ex- chambers for short periods, and analyses for ammonia, creted by the algae themselves. These findings are con- creatine, phosphate, and oxygen were made. A surprising sistent with the idea that during localized upwelling the result was that, under these conditions, ammonia and water comes from shallower levels and that during large- creatine were excreted in approximately equal propor- scale upwelling the water comes from considerably tions. It appears that creatine can be excreted in rather deeper levels. Water from deeper levels has not been in large quantities, and since it does not accumulate in sea- contact with the surface recently. An anodic-stripping water it follows that there is a very rapid turnover of this technique was used to directly measure trace metal con- substance. Phosphate appears to be excreted in a definite centrations and the effects of added chelators on trace ratio to creatine, and some preliminary calculations sug- metal concentrations in different water types. The results gest that large schools of anchovies moving slowly generally confirmed the hypothesis stated here. through an area may markedly influence the ammonia and phosphate concentrations in the water. A secondary effect of elevated ammonia concentration may be the re- Phytoplankton Ecology duced uptake of nitrate by phytoplankton. The research on phytoplankton ecology was under Urea in seawater was measured on the cruise. The an- the direction of a Spanish scientist and was carried out chovies used in the research just mentioned were also

41 43 monitored for urea excretion rates. The results suggested gorithrns have been developed and tested for stability that urea, like creatine, has a rapid turnover rate in the under different sampling conditions. They provide sam- sea. pling criteria that will be programmed into the shipboard computing syster:.. Benthos The benthic collections made on the Gosnold theand Thompson are being processed at the Woods Hole Ocean- SIMULATION MODEL PROJECT ographic Institution. The most interesting immediatere- Two submodels are being developedone for oceano- sult of these studies has been the discovery of largequan- tities of phosphorite in some of the collections obtained graphic processes and the other for nutrient cycling and in the vicinity of line 8 at 100 to 150 m (Figure 1). These biological productivity. deposits appear to form a pavement over the submarine A simulation model encompassing nutrient circulation, ridge that extends in a direction perpendicular to the phytoplankton production, and zooplankton grazing is shore. The relationship between the persistent upwelling being developed. This modeling effort providesa link in the vicinity mentioned and the presence of phospho- with the fisheries-oriented group in the Center for Quan- rite accompanied by high phosphate concentrations in titative Science at the University of Washington. the water is unknown. It is likely that these depositsare The oceanographic processes subrnodel is being devel- the source of anomalous phosphate concentrations found oped at Florida State University. The submarinetopog- in the water near the coast and over this ridge. Further raphy may be highly significant in determining regions of investigations of the phenomenon are desirable. potential upwelling. The Peruvian data should provide information useful in testing this possibility. In October 1969, plans were made to integrate the DATA HANDLING efforts in modeling of the oceanographicprocesses with the modeling of nutrient circulation and phytoplankton Data Center production. Virtually all the data taken during the cruise to Peru have been reduced and copies are available at the Depart- ment of Oceanography, University of Washington. A large AQUICULTURE PROJECT amount of data is included in a data bank on disk files and IBM cards. Lists of these data will be drawnup and A battery of three algal quasichemostatswere oper- circulated along with the instructions for requesting the ated on the roof of the oceanography building at the data from the data bank. Publication of data will be lim- University of Washington during the summer of 1969. ited to the routine hydrographic and productivity reports. Each was operated at a different dilution rate with water Plans are also being drawn up to maintain a complete obtained from Puget Sound and held in tanks. Seeded bibliography and reprint file on upwelling. This project with a plankton tow from Puget Sound, these chemostats will be computer oriented to allow rapid dissemination came to equilibrium rapidly, and initial populations com- of references to contributing investigators. posed mostly of dinoflagellates were displaced by diatom populations. As in the Friday Harbor chemostats of the Shipboard Computer Applications previous summer, the composition of the populations was a function of feed rate. The populations of the Fri- Two shipboard computer applications are being de- day Harbor experiment and of the Seattle experiments veloped. The first application is an integrated system for were similar in so many ways that investigators are con- data acquisition, editing, analysis, and presentation. The fident of success in their efforts to grow phytoplankton system provides for acquisition of up to 20 variables and with a relatively constant species composition and ata for graphical display of the distributions of the variables specified concentration to provide food for filter feeders. of interest. The system analyzes data from ..;'7D lowerings. Work on the aquiculture project will proceed at both The second application is use of the conTatingsystem the University of Washington and the Woods Hole Ocean- itself to make decisions about significant changes in the ographic Institution. Efforts at the University of Wash- phenomena under investigation. The system would then ington will continue to be directed toward the production trigger the appropriate sampling devices so that impor- of food phytoplankton of known species composition at tant events will not be missed and redundant data will controlled cell conc.entrations. These investigations will not be collected. be of a fundamental nature, concerned with the control Work is proceeding on a number of sampling algo- mechanisms that bring about steady state ina quasi- rithms that can be applied to the acquisition of data chemostat and with the conditions that permita multiple- while cruising or to sampling any general time series. Al- species population to exist. Plans for scaled-up versions

42 44 fr of these chemostats are being developed. An effort will 3. Lewin, University of Washington, Seattle (PS, Chaetocerus, silica) be made to utilize the effluent populations for feeding R. Mackenzie, Woods Ho lc Oceanographic Institution, Woods oysters and other filter feeders. Various mathematical Hole, Mass. (FC, benthic populations) models will be examined, and computer simulation tech- N. Margaris, University of Athens, Greece (PS, light limitation) M. McCarty, University of Washington, Seattle (computcr niques will be used where necessary. operator) J. McCarthy, Institute of Marine Research, University of California, San Diego (PS, urea studies) H. Minas, Station Marine d'Endoume, Marseille, France (FC, 15 N productivity) APPENDIX: PARTICIPANTS L. Olund, University of Washington, Seattle (computer programmer) On Board the R/V Thompson* R. Olund, University of Washington, Seattle (chemical analyst) R. Barber, Woods Hole Oceanographic Institution, Woods Hole, T. Packard, Univeisity of Washington, Seattle (PS, nitrate Mass. (PS, chclation effects on productivity) reductase, ammonia and phosphate regeneration) D. Blasco, Instituto de Investigaciones Pesqueras, Barcelona, M. Pamatmat, University of Washington, Seattle (NC, regenera- Spain (PS, phytoplankton taxonomy) tion of nutrients at sediment surface) N. Borovikoff, University of Washington, Seattle (marine D. Pillsbury, Oregon State University, Corvallis (PO, current technician) measurements) W. Broenkow, University of Washington, Seattle (NC, mathe- G. Rowe, Woods Hole Oceanographic Institution, Woods Hole, matical models) Mass. (FC, benthic populations) W. Chernoff, University of Washington, Seattle (chemical R. Smith, Oregon State University, Corvallis (PO, cutrent technician) measurements) J. Cline, University of Washington, Seattle (PO, formation of R. Stevenson, Bureau of Commercial Fisheries, National the oxygen minimum zone) Oceanographic and Atmospheric Administration, U.S. Depart- L. Conway, University of Washington, Seattle (FC, 15N ment of Commerce, Galveston, Tex. (PO, observer) productivity) C. Valardc, Hydrographic Service, Callao, Peru (PO, observer) C. Davis, University of Washington, Seattle (PS, quasi- V. Welsh, University of Washington, Seattle (marine technician) chemostate) T. Whit ledge, University of Washington, Seattle (FC, creatinc B. de Mendiola, Instituto del Mar del Peru, Callao, Peru excretion by anchovies) (FC, anchovy feeding habits) S. Zuta, Instituto de! Mar del Peru, Callao, Peru (PO, current R. Dugdale, University of Washington, Seattle (FC,15N measurements) productivity) J. Dugdale-Maclsaac, University of Washington, Seattle (PS, nutrient limitation) On Board the R/V Gosnold D. Enfield, Oregon State University, Corvallis (PO, current measurements) P. Clarner, Biological Oceanographer, Woods Hole Oceanographic R. Eppley, Institute of Marine Research, University of Cali- Institution, Woods Hole, Mass. (primary production studies) fornia, San Diego (PS, nitrate reductase and nitrate uptake) N. Corwin, Biological Oceanographer, Woods Hole Oceano- J. Goering, University of Alaska, College (NC, denitrification) graphic Institution, Woods Hole, Mass. (primary production 0. Guillen, Instituto del Mar del Peru, Callao, Peru (FC, 15N studies) productivity) B. Frost, University of Washington, Seattle (zooplankton P. Harrison, University of Washington, Seattle (PS, particulate sampling) silica) G. Grice, Woods Hole Oceanographic Institution, Woods Hole, M. Healy, University of Washington, Seattle (NC, trace metals) Mass. (zooplankton sampling) L. Hobson, Woods Hole Oceanographic Institution, Woods Hole, T. Lawson, Biological Oceanographer, Woods Hole Oceano- Mass. (NC, fate of particulate carbon) graphic Institution, Woods Hole, Mass. (zooplankton T. Hopkins, University of Washington, Seattle (PO, current sampling) measurements) Carl Lorenzen, Oceanography Section, National Science J. Kelley, University of Washington, Seattle (FC, mathematical Foundation, Washington, D.C. (phytoplankton studies) models, data processing programs) R. Mackenzie, Woods Hole Oceanographic Institution, Woods Hole, Mass. (phytoplankton studies) D. Menzel, Woods Hole Oceanographic Institution, Woods Hole, Mass. (primary production studies) G. Rowe, Woods Hole Oceanographic Institution, Woods Hole, *Key to abbreviations: PO = physical oceanography; NC = nu- Mass. (phytoplankton studies) trient circulation; PS = phytoplankton studies; FC = food J. Ryther, Woods Hole Oceanographic Institution, Woods Hole, chains. Mass. (primary production studies)

43 45 Marine Mammals

From antiquity to the present day, marine mammals drew up a proposed program in 1968. The National Com- have been a major resource through which man has uti- mittee and the International Marine Mammal Working lized the energy of marine ecosystems. While emphasis Group (PM) approved the proposal in 1969. on different species has changed through history, overall A recent proposal requesting support for thisprogram utilization has increased. With the growing protein and has beeri submitted..The Marine Mammal Working Group food shortages in the world, this trend can be expected (U.S.) has organized thisprogram. to continue. While studies of marine productivity have been focused on the base of the trophic structure, most of the exploitation has been at the top. This exploitation has, OBJECTIVES for the most part, been heedless of biological limitations, with the result that population after population has been depleted. This has been conspicuously true of marine The objectives of this program are to provide basic mammals. information needed for rational internationalmanage- Avoiding the repetition of past mistakes requiresmore ment of marine mammals and to encourage increased in- than good intentions, however; there must also exist ternational collaboration among marine mammalogists. sound biological information about a fauna within itseco- The program emphasizes the productivity of marinemam- mals within their trophic structure. Many marine system. Knowledge concerning marine mammals lagsa mam- century behind that concerning terrestrial mammals. This mals, especially the smaller species, ieflect unusually well program is an attempt to provide information about ma- the patterns of primary productivity of theseas since rine mammals on an ecosystem basis. they tend to concentrate where food resources are great- Traditional and current methods of study have empha- est. Further, because they must breathe air, these concentrations can be observed from the surface. sized population dynamics of economically valuedspe- cies, toe often with little ecological basis. Recent studies of behavior and physiology have emphasized the experimental animal, again with little opportunity foran eco- RESEARCH PLANS logical program. Only a vigorous and broadly based pattern of investigations by many persons working in a wide spectrum of CRUISES biological disciplines can yield an understanding that will be adequate for intelligent utilization and management of Since the distribution of ail marine organisms is highly the major biological resources that marine mammalsrep- discontinuous, it is important to study some of theareas resent. Studies of distribution, productivity, systematics, of discontinuity in detail. In Cetacea, discontinuities behavior, and ecology must go forward simultaneously if seem to be related to the levels of productivity of water understanding of marine mammals is to be obtained in masses. time to serve a useful purpose. Studies will have to beex- Cruises in the Bering Sea, in the Arctic Oceanadjacent panded to integrate investigations on and in thesea as to the Bering Sea, and in the South Atlantic Ocean (mid- reflections of species adaptations at many levels. The In- dle and high latitudes) are being planned. It is hoped that ternational Biological Program offersan opportunity for scientists from several nations will participate. marine mammalogists to cooperate in studies havingan ecological rather than an economic base. In 1967, the U.S. National Committee for the Inter- INFORMATION EXCHANGE national Biological Program decided that a study ofma- rine mammals should be included in the U.S./IBP and Publication of a newsletter for marine mammalogists appointed a group to develop a program. Thisgroup is expected to begin in 1971. It will includenews briefs,

44 46 notices of cruise opportunities, and items concerning G. Carleton Ray (Program Director), The Johns Hopkins recent literature. University, Baltimore, Md. International symposia are planned for further ex- Kenneih S. Norris, Oceanic Institute, Oahu, Hawaii change of information and to encourage research on ma- Charles 0. Handley, National Museum of Natural History, rine mammals. A symposium is being planned for the Smithsonian Insuiution, Washington, D.C. sacond year of this program to determine the status of William E. Schevill, Woods Hole Oceanographic Institu- marine mammal research and to stimulate greater partici- tion, Woods Hole, Mass. pation in the field. A second symposium is planned, for George A. Bartholomew, Jr., University of California, the final year of the program, to measure progress and Los Angeles establish directions for further research. These symposia John J. Burns, Alaska Department of Fish and Game, will be modeled after the First International Symposium Nome on Cetacean Research, held in Washington, D.C., in William C. Cummings, Naval Undersea Research and De- August 1963. velopment Center, San Diego, Calif. William E. Evans, Naval Undersea Research and Develop- ment Center, San Diego, Calif. Francis H. Fay, Arctic Health Research Center, College. STUDY CENTER Alaska Karl W. Kenyon, U.S. Fish and Wildlife Service, Seattle, A center for processing, storing and studying speci- Wash. mens of marine mammals is essential to progress in the Clayton E. Ray, National Museum of Natural History, program. Such a center is planned in Washington, D.C., Smithsonian Institution, Washington, D.C. by the Smithsonian Institution, which is participating in the program. Dr. G. Carleton Ray, Dr. Norris, Dr. Handley, and The center would include a cold room for temporary Mr. Schevill make up the Executive Committee of the storage of freshly captured specimens, a room for Council. Suzanne M. Contos is Executive Secretary. flensing and study, and equipment for macerating, de- greasing, drying, and embalming. It would also include equipment for preparing tissue for study under micro- scopes. RESEARCH METHODS Collections would inaa skeletons, soft parts preserved in fluid, frozi ,-,...arnens, and a variety of anatomical, histologic31, .ndpra trAogical specimens. The program is in three parts: Regional studies, including the relationships between cetacea and current systems of the Pacific, the distribution and abundance of South Atlantic cetacea as MANAGEMENT STRUCTURE correlated with primary productivity, and the biology of Bering Sea harbor seals and relationships with other ice- inhabiting pinnipeds The program is .F;ded by a Marine Mammal Council Studies of marine mammels of special interest, in- of 11 members, headed by a program director. The coun- cluding the social behavior of otariids as an influence on cil will develop the marine mammal study center, plan productivity and the reproductive behavior of the gray conferences and symposia, and coordinate various activi- whale ties, including field work; processing and storage of ma- New techniques, including population analyses of terials, aryl reporting to marine mammalogists. Council large cetacea through acoustics and the use of telemetry members are: for tracking and behavior studies of marine mammals

47 Origin and Structure of Ecosystems

The fundamental biological question that this program by studying the evolutionary history and the phyloge- is asking is whether two very similar physical environ- netic affinity of a few taxa within given phylogenetic ments acting on phylogenetically dissimilar organisms in groups different parts of the world will produce structurally and To determine the relatic.,ships of stability, diver- functionally similar ecosystems. If the answer is no, there sity, and productivity in these ecosystems cannot be any predictive science of ecology. In effect, To determine the relationships between native and knowledge acquired from studying a given ecosystem can- exotic plants and animals, and man's role in the introduc- not be applied to an anologous ecosystem, unless similar tion and establishment of weedy organisms physical environment indeed means similar ecosystem. To establish the interrelationship between ecosys- This program has three components. One is a study of tem structure and human culture and economy ecosystems that have evolved separately under similar physical environments to ascertain whether they have similar structure and ecological niches. Another is a study of evolutionary and ecological diversity in selected groups RESEARCH PLANS of organisms, particularly amphibians of the New World. The third component is a study of an isolated island ecosystem (Hawaii). It will be concerned with the diversity and stability of isolated ecosystems and with the effect STRUCTURE OF ECOSYSTEMS of isolation on structure. Common to these three subprograms is the intent to The research strategy for this subprogram is to com- integrate studies of ecology with studies of evolution to pare two sites for each biome. The biomes to be consid- investigate ecosystem structure and development. Eco- ered are the thorn scrub and Mediterranean scrub. One logical aspects considered are species diversity, niche uti- biome site is in South America; the other is in North lization, and life form interaction. Evolutionary aspects America. considered are the causes, mechanisms, and rates of For the thorn scrub vegetation, comparisons are being speciation. made between the Silver Bell Bajada site, near Tucson, The research in this program requires study of ecosys-. Arizona, and one near Andalgala in the state of Cata- tems in widely separated areas. Hence, this program has marca, Argentina. FM' the Mediterranean scrub, compari- developed as an international effort involving scientists in sons are beiag made between Cuesta la Dormida, prov- many American countries. ince of Santiago, Chile, and Corte Madera Valley of the Laguna Mountains, California. Field and laboratory work is designed to compare the environments and to compare the ecosystems. Elements CBJECTIVES of the environment to be considered are macroclimatic and microclimatic patterns. soil genesis (including soil micromorphology), geological history and paleohistory, Objectives are: and land-use patt- .ns. Elements of the ecosystems to be To investigate ecological systems with similar physi- considered are structure (including physiognomy, mor- cal environments and determine the degree of ecological phology, species number and diversity, life forms, niche relationship between the species patterns, and pattern of distribution of individuals in the To discov-r how much of this similarity is attribut- community), floristic and faunistie similarities and dif- able to parallel evolution of relatively closely related taxa ferences, and function (including phenological cycles. and how much is attributable to convergence of distantly regulatory mechanisms, succession patterns, energetic related taxa and biochemical pathways, adaptive modes, and species To improve understanding of ecological diversity interactions).

^48 EVOLUTIONARY AND ECOLOGICAL Ocean (Hawaiian Islands). These ecosystems contain the DIVERSITY same principal environmental componentsthat are found in many mainland tropical ecosystems, but their biotic original The objective of this subprogram is to improveunder- component is extremely restricted in origin. This standing of the evolutionary history and ecological di- biotic component has evolved into a diversity of its own. versity of a few taxa. The approach is multinational and To the endemic diversity of the island ecosystems a non- multidisciplinary. Anuran amphibians of the New World evolutionary diversity has been added through man- are being studied by cooperating teamsof North and introduced biota. A competitive struggle is going on in endemic South American workers. many ecosystems that threatens to destroy the Anuran amphibians are most diverse in the tropics, diversity in several of the island habitats. This subprogram seeks to discover the fundamental and little ecological or taxonomic informationabout gain in- them is available. Consequently, much fundamental eco- causes of the competitive struggle in order to logical and taxonomic research is necessary. sights into the stability-fragility relations of those native been Two types of studies will be conducted: ecological ecosystems where the direct interference of man has and phylogewnic. Intensive ecological studies inecologi- held to a minimum. This subprogram consists of intensive studies of a few cally different areas will include measurements oflarval and adult microdistribution, measurements of speciesdi- selected ecosystems, comparisons of ecosystems along versity, seasonality, trophic position, population qizeand well-defined environmental gradients, and studies of en- mechanisms, and demic populations. The ecosystems studied intensively struc tu re, reproductive cycles, isolating and structure have niche breadth. The standardized techniques used inthese are those whose species composition studies will be comparable with those used in recent been the result of island evolution. The intensive ecosys- and struc- major studies in the Old World tropics. tem studies are to determine the composition Neotropical anuran amphibians are being studied with tural variation of the ecosystem, the direction of the respect to external morphological features ofadults and ecosystem's development, and the rates of population tadpoles used in taxonomy; osteological characters and change. their importance in the determination of supraspecific re- Ecosystems are compared along environmental gra- lationships; sperm morphology; karyotype, including dients in order to study the factors responsible for number and structure of chromosomes; biochemical fea- changes in ecosystem stability. These studies will investi- tures of skin, blood, and other systems; life histories; gate the nature of the relationship betweenspecies di- premating isolating mechanisms, especially mating calls; versity and ecosystem stability. The interrelationship of and experimental hybridization. ecosystem structure, dominant organisms, and ecosystem The phylogenetic and ecological studies complement structure along an environmental gradient willalso be each other. Many data of interest to the group concerned investigated. with phylogeny will derive from the site studies, and the Endemic populations that successfully adapted to phylogenetic group will provide the ecologically oriented island ecosystems will be studied to determine the mech- group with estimates of phyletic distancebetween the anism of adaptation. Specific points to be considered are animals they are studying. the factors that determine success, rates of speciation, The major reasons for emphasizing frogs (order Anura) and degree of speciation. in the study are as follows: Because of their abundance and their important position in the food chain, frogs are a highly significant MANAGEMENT group in many ecosystems. Frogs are easily collected and easily maintained in This program is directed by W. Frank Blair. The organizational structure is shown in the following list: the laboratory. A multidisciplinary approach to anuran research is possible because of the wide variety of techniques that Structure of ecosystems (Otto T. Solbrig) T. have been found successful, and this approach greatly in- Desert scrub project (Jorge Morello and Otto creases the reliability of phylogeneticconclusions. Solbrig) Mediterranean scrub project (Francesco di Castriand Harold Mooney) ISLAND ECOSYSTEM STABILITY Evolutionary and ecological diversity (Craig E.Nelson AND_EVOLUTION and Sheldon Guttman) Island ecosystem stability and evolution(Andrew J. This subprogram is concerned with tropical ecosys- Berger, J. Linsley Gressitt, and Dieter Mueller- tems that evolved in isolation in the midst of thePacific Dombois)

47 49 RESEARCH METHODS AND PROGRESS the comparable North American area in havinga depau- perate fish fauna. STRUCTURE OF ECOSYSTEMS Working on the population genetics of Bufoarenarum, Terry Matthews collected breeding stocks during his field Desert Scrub work in Argentina in the fall of 1968 and from these did laboratory crosses to learn the genetic basis for the be- During early 1971, studies will begin at the sitenear havioral trait he is studying in wild populations. He did Andalgald. In late 1970, transects will be set up anda additional field work from September to December1969. base map, describing the vegetation, will be prepared. In 1969, John Kirsch and an Australian coworker started Several preliminary studies have been made of the several months of collecting marsupial material in north- biota of the semidesert areas of Argentina. These projects, ern and western South America. In the fall of 1969, Jerry which contribute to the work at Andalgala, area remote Boggs, graduate student supervised by Don Hunsaker, be- sensing flight and studies of subandean insects, animal gan a population study of Marmosa and Caluromys in niches, genetics of Bufo arenarum, and genetics of thorn Minca, Colombia, in the Santa Marta Mountains, where scrub. living research facilities were provided by the Instituto Previous work by Morello with vegetation of the Ar- de Recursos Naturales, an agency of the Colombiangov- gentine Monte provides a starting point for the analysis ernment. of the Argentine component of the two convergent thorn In 1969, research on the genera Larrea and Glandu- scrub systems being compared. A remote sensing flight laria, and work on Prosopis, was initiated by OttoT. by the National Aeronautics and Space Administration Solbrig of Harvard University and Juan H. Hunziker of (Earth Resources Division Mission No. 97)over northern the University of Buenos Aires. Argentina from Formosa to Salta on July 19, 1969,pro- The first phase of the program of crosses between vided invaluable data for the vegetational aspect of the North and South American species of Glandularia has study. The flight was a cooperative effort of NASA and been completed and some of the results have been pub- of personnel of the Instituto Nacional de Tecnologia lished. It was demonstrated that the NorthAmerican Agropecuaria (INTA). Instrumentation included two hexaploid species G. elegans is a segmental all ohexaploid RC18 mapping cameras (Ektachrome MA Aerographic of diploid South American species Working indepen- and Ektachrome IR Aerographic film), one multiband dently and using cultivated species, Khoshoo in Indiawas KA62 camera system (three images with Plus-S Aero- able to arrive at similar results in regard to the North graphic film at wavelengths of 477, 530, and 517),one American hexaploid G. canadensis, which he crossed with camera system with infrared film at 750, and a thermal the cultivated G. hybrida, a species of South American two-channel mapper. An Auxiliary Data Annotation Sys- origin. Crosses between several North American species tem provided all necessary information about date, time, by Solbrig, including G. elegans and G. canadensis,indi- ahitude, ground speed, and the like. cate that they are closely related cytologically. It is there- INTA personnel are responsible for "ground truth" fore valid to hypothesize that the North Americanspe- for the remote sensing data in Argentina and for interpre- cies are derived from South Americanancestors. Whether tation. Complete sets of the data collected in Argentina the North American species originated in South America, have been provided by NASA to the INTA organization migrated northward, and later became extinct in the in Argentina and to the University of Texas in Austin. south, or whether the South American speciesonce grew Field work leading to a comparison of ecological in North America and there gave rise to the polyploid niches of animals is most advanced for lizards and fresh- species, is impossible to tell. The cituation in Glandularia water fishes. Richard D. Sage has completed 3 years of appears to be unique, since it cannot be explained bya field work in western Argentina and has identifieda se- single migration event as in thecase of other disjunct dis- ries of ecological equivalents of North American species tributions involving interfertile plants. among the lizards. He has concentrated on detailed stud- The genus Larrea is represented in South Americaby ies of the eLological life history of a few species (Li- four species, three of which are shrubs Ito 3 m high: olac darwini = Uta stansburiana and Phymaturus L. cuneifolia, L. divaricata, and L. nitida. Theyare dis- palluma = Sauromalus obesus) but has also workedex- tributed from Peru to Chubut and from Chileto Buenos tensively with other species. William S. Birkhead, who Aires. The fourth species is a woody chamaephyte (L. terminated a year and a half of field work in early 1970, ameghinoi), whose range extends in Patagonia froca has been working with the freshwater fishes of the Neuquen to Chubut. streams that drain from the Sierra Cordoba. Preliminary Cytological studies done by Juan H. Hunziker have results reveal striking similarities in the ecological niches shown that L. divaricata, L. nitida, and L. ameghinoiare of these fishes and those of the comparable North Ameri- diploid (n=13), whereas L. cuneifolia isa tetraploid can area. Streams at the Cafayate site resemble those of (n=26).

50 It was possible to locate and study in detail areas of 196i-1968 in the upper Amazon Basin of Peru col- where all four species meet. Hunziker et al,(1969)* lecting karyotype material, making hybridization and found that in such situations hybrid swarms are formed ecological studies, and recording mating calls. He col- from hybridization between L. ameghinoi and L. nitida lected similar material in Colombia in November 1968, on one hand and L. nitida and L.cuneifolia on the other. and in Argentina and Brazil from September to Deceni- Studies of seed protein and flavonoid patterns have been ber 1969. In Brazil he obtained karyotype material for performed or are in progress on all the South American 75 species representing 26 genera and 7 families of species and on the various ecotypic forms of the North anurans. American species, L. tridentata. Several undescribed species were collected in the Vuilleumier (1969)* has written a biosystematically course of the Peruvian work. One ancillaryaccomplish- oriented revision of the South American species of the ment was to ascertain that two speciesof Bufo have been genus Perezia, a genus with a disjunct NorthAmerican- confused unckr the name B. marinus. Both exist sympat- South American distribution. rically at Iparia. B. poeppigi is relatively small and slender In January 968, the entomology section of the Insti- and has narrow parotid glands; B. rnarinus is large and ro- tuto Miguel Lilo, in Argentina, initiated a 5-year study bust and has large parotid glands. These species differ of the insects of the subandean biota (Southern Creosote karyologically, in mating call, and in crossability with Bush Province). Financing of this project is shared by the other species of Bufb. Instituto Miguel Lillo and the Consejo Nacional de In- M. J. Fouquette collected Hyla and recorded mating vestigaciones Cientificas y Tecnicas. The principal investi- calls in Colombia and Venezuela from November toDe- gators are Abraham Willink (Hymenoptera),Lionel A. cember 1967 and in Panama and Colombia during the Stange (Neuroptera), and Arturo Teran (Coleoptera: summer of 1969. Papers in preparation as aresult of this Bruchidae). work include: "Mating Calls of Colombian Frogs," by The first phase of t.he entomological study, which is M. J. Fouquette, William F. Pyburn, and V. H.Hutchison; concerned with making general collections, began in "Review of the Frogs Related to Hyla boulengeri,""The Note 1968 andscheduled for 2 years. Ten malaise traps were Identity of Hyla rubra and Related Species," "Brief installed in the first year, and samples are coileotez on Hyla rufitela," "BreedingCalls of Frogs in Guyana 2-week intervals. Regular collecting and study trips into and Surinam," and "Hyla microcephala and Related the area are being made, and a mobile laboratory is used Frogs in Northern South America," by M. J. Fouquette; to facilitate studies in remote areas. In the second phase and "Morphology and Evolutionary Trends inAnuran of the study, scheduled for 3 years, more specialized Spermatozoa," by A. James Delahoussaye. studies will be made. Studies already under way are con- William F. Pyburn and a student assistant spent June :alls and cerned with bee pollinators of biogeographically oi eco- and July in eastern Colombia recording mating logically import:int plants; insects associated with Larrea; collecting stocks of Hylidae. Richard Newcomer, post- seed-feeding insects, especially Bruchid beetles; and sand doctoral investigator, spent 2 months in the Choco prov- and 3 months dune communities, especially the Myreloontidae. ince of northeastern Colombia in late 1968 in the Ilanos of eastern Colombia in early 1969collecting mating calls and experimental stocks ofHylidae. He re- Mediterranean Scrub turned to Colombia in September 1969 to help establish Detailed plans were made for work on the convergence a biochemical laboratoryfor the amphibian work at Uui- of this system, as represented in California and Chile.Re- versidad Javeriana, Bogota, where Jaime George is co- search will begin as soon as funds become available. operating in the project. Luis Dino Vizotto, University São Jose doRio Preto, Brazil, spent 2 months at the University of Texasin early EVOLUTIONARY AND ECOLOGICAL 1969 to learn techniques of biochemical studies, karyo- DIVEIRSITY logical studies, and hybridization. In October 1969, William F. Martin, Ford Foundation Preliminary efforts have been toward understanding postdoctoral fellow, went to Medellin, Colombia,where the evolutionary relations of the huge tree frogfamily he is establishing a bioacoustical center for theamphibian Hylidae. The scope of this subprogram has been enlarged research. to include New World anurans, although the families W. Frank Blair collected s:;xperimental stocks of am- Hylidae and Leptodactylidae will receive high priority. phibians in Colombia and Argentina in January 1969, Several field expeditions have been made. James P. and in early February he ccllected with AlbertoVeloso Bogart, postdoctoral investigator, spent the rainy season ofValparaiso and Klaus Busse of Santiago in southern Chile. All six species of the leptodactylid frogs ofthe *See Appendix A (p. 50). genus Eupsophus were collectedalong with representa-

49 tives of various other taxa. Subsequent analysis of the logical and two climatic stations were established in dif- Eupsophus karyotypes by James Bogart revealed unex- ferent macroclimates along the transects. pected variability: four different chromosome numbers. Evolution Studies on Endemic Populations ISLAND ECOSYSTEM STABI LITY Most of this work has been done along the transects, AND EVOLUTION but some highly specialized populations, suchas the Ha- waiian Diptera and certain Hawaiian birds, have been Intensive Study of Selected Ecosystems studied in different areas. Thr same is true for thegene- cological studies of the widely distributed polymorphic A comprehensive ecological survey was begun ina tree Metrosideros. The evolutionary objectives are pur- 200-acre site occupied by a native montane foresteco- sued on all major islands, particularly on Hawaii, Maui, system. The principal structural components are scat- and Kauai. They are not necessarily tied to the transects. tered, tall, big-diameter Acacia koa trees and groups of The ecological studies will be extended eventuallyto Metrosideros polymorpha with several other nativetree Mani and Kauai. species as subcanopy members. A third major biomass stratum is formed by tree ferns (Cibotium spp.). The base lines were surveyed and two 1,000-m-long transects were flagged out with 10 plot points at intervals APPENDIX A: PUBLICATIONS of 200 m. The first sample plot has been completed. In Auffenberg, W., and W. G. Weaver, Jr. 1969. Gopherus addition to the plant ecological survey, insect and bird berlanderi in Southeastern Texa,. Bull. Florida State Museum 13. 3 p. surveys are carried out along the same transects. Malaise Blair, W. F.1969. Especiación en los sapos (genero Bufo). traps and attractant logs have been placed at various stra- Acta Zool. Lilloana XXIV, p. 317. tegic positions in this forest. Blair, W. F. 1970. The influence of the Amazon River Basinon the evolution of toads of the genus Bufo of South America. A climatic station was installed in August 1970 for II Simposio y Foro de Biologia Tropical Amazonica: 168- continuous recording of temperature and humidity. Rain- 171. fall in an open area and under trees is checked at weekly Fouquette, M. J., Jr. 1968. Some frogs from the Venezuelan Ilanos, and the status of Hyla misera Werner. Herpetologica intervals. Precipitation measurements under thecanopy 24: 321-325. are particularly important here, because of the suspected Hunziker, J. H., R. A. Palacios, and A. Soriano. 1969. Hibrida- high moisture contribution from cloud interception. cion Natural en especies sud americanas de Larrea (Zygo- phYllaceae). Kurtziana 5:55-56. Checklists of all biota in this forest have beenpre- Solbrig, 0. T., C. Passar,, and R. Glass. 1968. Artificial pared, and further sampling techniques are being worked hybridization between different ptdloid levels in Glandu- out. laria (Verbenaceae). Amer. J. Bot. 55:1235-1239. Vuilleumier, H. S. 1969. Evoluticn and systematics of Perezia sect. Perezia (Compositae-Mutisieae). Contr. Gray Herb. 199. Ecosystem Transect Study Six ecosystem transects have been defined and described on the island of Hawaii. Theserange from 10 to APPENDIX B: PARTICIPANTS* 22 miles in length. One extends from the top of Mauna W. Frank Blair, Department of Zoology, University of Texas, Kea (elevation about 14,000 ft) to the sugarcane fields Austin (Program Director; SE, niches and microcommuni- above Hilo. Another lies parallel to the Mauna Keatran- ties; EED, anuran work) sect on the east flank of Mauna Loa, where it traverses Homer Aschman, Department of Geography, University of California, Riverside (SE, land use) similar climates but much younger substrates. The four Peter D. Ashlock, Department of Entomology, Bernice P. remaining transec is are in Hawaii Volcanoes National Bishop Museum, Honolulu, Hawaii (IESE, phytophagous Park and extend to the Mauna Loa transect down tosea insects, Heteroptera) Geoffrey C. Ashton, Department of Genetics, University of level. Hawaii, Honolulu (IESE, genetic analysis of populations) Each of the 30 principal ecosystems along thesetran- Franceso J. Ayala, Rockefeller University, New York, N.Y. sects has been studied in two 500-m2-plots. The plot lo- (BED, Drosophila) Gladys E. Baker, Departmelit of Botany, University of cations are being transferred to a new vegetationmap Hawaii, Honolulu (IESE, ecological roles of the fungi) having a scale of 1:24,000. Field work for themap has Michael G. Barbour, Department of Botany, University of been completed. Sixty-two plant checklists containing California, Davis (SE, vegetational structure) John W. Beardsley, Department of Entomology, University of quantitative information on species coverare pro- Hawaii, Honolulu (IESE, the effects of sap-sucking Homop- grammed for computer storage and retrieval. Various tera on the stability and fragility of Hawaiian ecosystems) summaries of this information will be available forany project participant. The same procedure will be followed "-The abbreviation; SE, IESE, and EED refer to subprograms. SE = structure of ecosystems; IESEisland ecosystem stability for checklists of insects and other biota. Seven pheno- and evolution; EED = evolutionary and ecological diversity.

50 52 Andrew J. Berger, Department of Zoology, University of Don Hunsaker, Department of Zoology, San Diego State Col- Hawaii, Honolulu (Co-Director, IESE; life history and func- lege, San Diego, Calif. (SE, mammal niches) tional anatomical studies of the Hawaiian honeycreepers) Juan H. Hunziker, Department of Genetics, University of William S. Birkhead, Department of Zoology, University of Buenos Aires, Argentina (SE, larrea and prosopis genetics and Texas, Austin (SE, fish niches) biochemistry) James P. Bogart, Department of Zoology, University of Texas, Jaime Hurtubia, Instituto de Ecologia, Universidad Austral de Austin (EED, anuran work) Chile, Valdivia (SE, leaf insect niches) Jerry Boggs, Department of Zoology, San Diego State College, Albert W. Johnson, Dean of Sciences, San Diego State College, San Diego, Calif. (SE, mammal niches) San Diego, Calif. (SE, plant species diversity) Charles D. Bonhani, Department of Watershed Management, John M. Kingsolver, Agricultural Research Service, U.S. Depart- University of Arizona, Tucson (SE, vegetational structure) ment of Agriculture, Washington, D.C. (SE, arthropod D. E. Bradbury, Department of Geography, University of Cali- niches) fornia, Los Angeles (SE, geomorphology) John Kirsch, Department of Zoology, University of Kansas, Nadir Brum-Zorilla, Institu to de Biologia, Montevideo, Uruguay Lawrence (SE, marsupial convergence) (EED, anuran work) Charles H. Lamoureux, Department of Botany University of Klaus Busse, Centro de Investigaciones Zoologicas, Santiago, Hawaii, Honolulu (IESE, growth rates and anatomy of woody Chile (EED, anuran work) plants in relation to environment) Martin Cody, Deparanent of Zoology, University of California, Robert M. Lloyd, Department of Botany, University of Hawaii, Los Angeles (SE, bird niches) Honolulu (IESE, life history studies of ferns) Carolyn Corn, Department of Zoology, University of Hawaii, Charles H. Lowe, Department of Zoology, University of Ari- Honolulu (IESE, studies of Metrosideeros) zona, Tucson (SE, vegetational structure) Chieita Culbersor Department of Botany, Duke University, Bertha Lutz, Museu Nacional, Rio de Janeiro, Brazil (EED, Durham, N.C..(SE, lichen ecology) anuran work) William Culberson, Department of Botany, Duke University, Tom J. Mabry, Department of Botany, University o, Texas, Durham, N.C. (SE, lichen ecology) Austin (SE, plant biochemistry) Clifton J. Davis, Chief Entomologist, Hawaii Department of Michael A. Mares, Department of Biology, University of Texas, Agriculture, Honolulu (IESE, Hawaiian Cerambycid beetles Austin (SE, rodent niches) and ceitain other wood borers) William F. Martin, Universidad de Antioquia, Medellin, Colombia A. James Delahoussaye, Department of Zoology, Arizona State (EED, anuran work) University, Ternpe (EED, anuran work) Terry Matthews, Department of Zoology University of Texas, Mercedes D. Delfinado, Department of Entomology, University Austin (SE, amphibians) of Hawaii, Honolulu (IESE, biosystematics of Hawaiian Ming-Pi Mi, Department of Genetics, University of Hawaii, Diptera) Honolulu (IESE, computerization) Hernando de Macedo, Universidad Cayetano, Lima, Peru Phillip C. Miller, Department of Biology, San Diego State Col- (EED, anuran work) lege, San Diego, Calif. (SE, microclimate) Reinaldo Diaz, Department of Zoology, University of Texas, Wallace C. Mitchell, Department of Agriculture, University of Austin (EED, anuran work) Hawaii, Honolulu (IESE, biology-ecology and control of in- Francesco di Castri, lnstituto de Ecologia, Universidad Austrai sects attacking the silverswords in Hawaii) de Chile, Valdivia (Co-Director, Mediterranean Scrub Pro- Andrew R. Moldenke, Department of Biology, Stanford Univer- ject, SE; soil conununity structure) sity, Stanford, Calif. (SE, pollination systems) Theodosius Dobzhansky, Department of Zoology. Rockefeller Harold Mooney, Department of Biological Sciences, Stanford University, NeW York, N.Y. (EED, Drosophila work) University, Stanford, Calif. (Co-Director, Mediterranean Maxwell S. Doty, Department of BotanY, University of Hawaii, Scrub Project, SF; vegetational structure) Honolulu (IESE, ecolelical and evolutionary roles of the Jorge Morello, Instituto Nacional de Tecnologia Agropecuaria, algae) Buenos Aires, Argentina (Co-Director, Desert Scrub Project, Lee Ehrman, Department of Population Genetics, Rockefeller SE; vegetational structure) University, New York, N.Y. (EED, Drosophila work) Dieter Mueller-Dombois, Department of Botany, University of M. J. Fouquette, Department of Biology, Arizona State Uni- Hawaii, Honolulu (Co-Director, IESE; vegetation-environ- versity, Tempe (EED, anuran work) ment correlation studies, studies of distributional dynamics, Wayne C. Gagne, Bernice P. Bishop Museum, Honolulu, Hawaii life history studies of important plants: angiosperms) (IESE, phytophagous insects, Heteroptera) Craig E. Nelson, Department of Zoology, Indiana University, Jaime George, Universidad Javeriana, Bogota, Colombia (EED, Bloomington (Co-Director, EED; anuran work) anuran work) Richard Newcomer, Department of Biology, Univc.,sity of Texas, Roger D. Goos, Department of Botany, University of Hawaii, Austin (EED, anuran work) Honolulu (IESE, ecological roles of the fungi) Toshiyuki Nishida, Department of Entomology, University of J. Linsley Gressitt, Department of Entomology, Bernice P. Hawaii, Honolulu (IESE, behavior and population dynamics Bishop Museum, Honolulu, Hawaii (Co-Director, IESE; of endemic insects) Hawaiian Cerambycid beetles and certain other wood Dan Otte, Department of Zoology, University of Texas, Austin borers) (SE, arthropod niches) Sheldon Guttman, Miami University, Oxford, Ohio (Co- William F. Pyburn, Department of Biology, University of Texas, Director, EED; anuran work) Arlington (EED, anuran work) Ernst R. Hajek, Department of Ecology, Universidad Catolica Frank J. Radovsky, Acarologist, Bernice P. Bishop Museum, de Chile, Santiago (SE, microclirnate) Honolulu, Hawaii (IESE, parasites of vertebrate animals of Frank H. Haramo to, Department of Entomology, University of Hawaii) Hawaii, Honolulu (IESE, behavior and population dynamics Dennis Ralin, Department of 'Lc ology, University of Texas, of endemic insects) Austin (EED, anuran work) D. Elmo Hardy, Department of Entomology, University of Peter Raven, Department of Biology, Stanford University, Stan- Hawaii, Honolulu (IESE, evolution and genetics of Hawaiian ford, Calif. (SE, pollination systems) Drosophilidae, biosystematics of Hawaiian Diptera) Juan Rivero, University of Puerto Rico, Mayaguez (EED, anuran Fabio Heredia, Universidad de Antioquia, Medellin, Colombia work) (EED, anuran work) Herbert H. Ross, Entomology Department, University of Jorge Hernandez, Instituto de Ciencias Naturales, Bogota, Colom- Georgia, Athens (SE, arthropod niches) bia (EED, anuran work)

5 1 Pedro M. Ruiz-C., 1Jnivcrsidad Nacional de Colombia, Bogota, Hawaii, Honolulu (IESE, effect of diseases in endemic popu- Colombia (EED, anuran work) lations of insects) Francisco Alberto Sacz, lnstituto de Biologia, Montevideo, Arturo Teran, Department of Zoology, Instituto Miguel Li llo, Uruguay (EED, anuran work) Sat: Miguel de Tucuman, Argentina (SE, insects) Richard D. Sage, Dcpartment of Zoology, University of Texas, Wallace B. Thompson, Departincnt of Biology, University of Austin (SE, lizard niches) Texas, Austin (SE, plant biochemistry) Francisco Saiz, Department of Entomology, Universidad Austral Norman J. W. Thrower, Department of Geography, University de Chile, Valdivia (SE, beetic niches) of Californ; os Angeles (SE, geomorphology) G. Allen Samuelson, Departmcnt of Entomology, Bernice P. P. Quentin .lich, Hawaii Department of Health, Honolulu. Bishop Museum, Honolulu, Hawaii (IESE, Hawaiian Ceram- (IESE, pops. ation structure of rodcnts, brecding and feeding bycid beetles) patterns uf feral goats and pigs) Eduardo Sanchez, Universidad Nacional de Chi lc, Santiago (SE, B. L. Turncr, Dcpartment of Botany, University of Texas. plant biochemistry) Austin (SE, plant biosystematics) Peter Seligmann, Bioquimica, Instituto Miguel Lillo, San Migucl Alberto Veloso, Universidia! de Chile, Valparaiso (EED, anuran de Tucuman, Argentina (SE, plant biochemistry) work) Richard B. Se !ander, Department of Entomology, University of Valeria Vitali di Castri, Universidad Au Ara.i de Chile, Valdivia Illinois, Urbana (SE, arthropod niches) (SE, arachnid niches) Jean-Pierre Simon, Department of Botany, Claremont Graduatc Luis D'no Univcrsidad dc STo Jose do Rio Prao, Brazil School, Claremont, Calif. (SE, plant biochemistry) (EED, anuran work) Otto T. Solbrig, Department of Biology, Harvard University, Beryl S. Vuilleumier, Gray Herbarium, Harvard University, Cam- Cambridge, Mass. (Director, SE; Co-Director, Desert Scrub bridge, Mass. (SE, plant biosystematics) Project, SE; plant genetics and biochemistry) Gordon D. Wallace, Departmcnt or Tropical Medicine, National Lionel A. Stange, Department of Zoology, Instituto Migue: Institutes of Health, Queen's Hospital, Honolulu (1ESE,.. Lillo, San Migucl dc Tucuman, Argentina (SE, insects) parasites of vertcbrate animals in Hawaii) Wallace A. Steffan, Department of Entomology, Bernice P. Abraham Willink, Departmcnt of Zoology, lnstituto Miguel Bishop Museum, Honolulu, Hawaii (IESE, ecology and Li llo, San Miguel de Tucuman, Argentina (SE, insects) cvolution of Hawaiian Sciaridae) T. Wei Yang, Arizona Sonora Desert Museum, University of Minoru Tamashiro, Department of Entomology, University of Arizona, Tucson (SE, Larrea communities)

52 HUMAN ADAPTABILITY COMPONENT Popuiation Genetics of South American Ind ians

Human adaptability has two components: the genetic, turated tribes will provide a critical base line for our un- which sets the limit of the organism's ability to respond, derstanding of human adaptability to a wider, more com- and the physiological, which determines the range ofie- plex range of social and technological circumstances. sponse available to the organism within the genetic limit. Man's genetic i:iversity and most of his adaptations With the development and ascendancy of agriculture arose while he was living at the tribal level of culture, as some 10,000 years ago in the ad. World, and even more a hunter and gatherer or as a neolithic farmer. There are recently in the New, one of man's most successful social many insights into the population genetics of civilized forms, the tribe, was transformed successively into the man that can be gained only from studies on such primi- city, the state, and, ultimately, the nation. Dramatic tive populations. The term "primitive" as here used changes also took place in the realm of technology and means a population socially and politically organized demography, increasing both man's need and his ability along lines of kinship, the hallmark of tribal society. Ac- to penetrate new ecological zones on an unprecedented tually. many aspects of "primitive" society are breath- scale. These dramatic changes took place within the last taking in their sophistication and complexity. The resil- percrnt of human cultural and biological evolution. ience and durability of many aspects of tribal social One of the consequences is that the present major popu- organization are evidenced in the many rural, peasant lations of the world owe their numerical and biological papulations of the world today. signincance to very recent events. This program is directed by James V. Neel, Chairman, It_ his long career as a tribesman, man adapted, both Department of Human Genetics, University of Michigan culturally and physically, to a wide range of climates and Medical School, Ann Arbor. geographical conditions. The Age of Discovery, barely a few centuries behind us, found tribal cultures successfully and intimately adapted to boreal and tropical forests, tundra, deserts at high and low altitudes, grasslands, the OBJECTIVES arctic, and insular, riparian, and riverine ecological zones. Wherever such cultures were found, they exhibited a remarkable variability in specific technological adjustments This program seeks to answer four basic questions of required by particular environments. Socially and demo- scientific interest: graphically there was, however, a remarkable degree of What is the tempo of human evolution as measured similarity among tribesmen. by the degree of divergence that has arisen between In- The recent and continuously accelerating spread of dian groups since their arrival in the Americas? civilized culture has replaced or transformed many of the For such Indian groups as still exist in a pre- adaptations that are known to have existed, but there Columbian state, what significant biological parameters still remain a number oc very significant tribal popula- affected their evolution? tions in various parts ol the world. Some of these can be What patterns of social and biological adaptation taken to approximate th. social, demograrhic, epidemi- emerge as primitive Indian groups are subjected to cul- ological, and ecological circurnstancc:s un,ler which man tural change? evolved for the greater par of his history. To what extent do cultural and social practices Because of man's long history as a tribesman an I the (marriage, settlement pattern, intergroup hostility) af- surprisingly narrow ral,e of social types exhibited fect the biological and demographic characteristics of among tribesmen, the study of these remaining unaccul- the population?

55 RESEARCH METHODS AND PROGRESS a consequence of the fact that the Vanonmma periodically raid the Makiritare and abduct women from them. In one instance a unique and previously unreported gene To obtain the basic data necessary to answer these (a serum albumin gene) was discovered in a Yanomama questions, specialists from several disciplines participate village in an abducted Makirita re tribeswoman and her in both the actual field studies and the laboratory analy- progeny, she being the only survivor of her tribe. The ses of the biological specimens. These disciplines include others were exterminated by the Yanornama (Weitkamp social anthropology, physical anthropology, linguistics, and Chagnon, 1968; Chagnonet al.,1970). medicine, genetics, serology, and chemistry. Our partici- We have taken some 30,000 ft of film on the Yano- pants in Latin America include members of the Venezu- mama. This has been obtained both as a research re- elan IBP and the Brazilian IBP. source and for preparing several teaching films. The The principal efforts of this program are directed to- teaching films are needed because of the difficulty e> ward the Yanomama Indians or Venezuela and Brazil. perienced in describing the nature of the Yanomama to Less extensive studies have been performed on the a student or ccientifie audience. One ethnographic film Xavante of Brazil and the Makiritare of Venezuela and "The Feast," has already been prepared.* It depicts a Brazil. The Yanomama speak only their nitive language, central social and political function in the political life of an American Indian tongue that is thus far not demon- the Yanornama. Two films are in preparation. strably related to any of the well-described linguistic groupings of South America. Accordingly, it was necessary from the outset to develop the anthropological and GENERAL HEALTH AND REPRODUCTION linguistic groundwork on which the feasibility of many of the related studies depended. The cultural anthropolo- The general health of the Xavante and Yanomarna has gist spent 15 months with the tribe prior to the initiation been assayed by vital statistics and la; physical examina- of the medical and genetics work. During this time he actions on an unselected cross section. The vital statistics quired a proficiency in the language and established rap- on the Xavante have been summarized (Salzanoet al., port and contact with the villages that were later seiected 1967). The more extensive material on the Yanomama for study. Genealogical and demographic data and infor- has been published in preliminary form (Neel and mation on recent village histories and interrelationships Chagnon, 1968). An accurate demographic characteriza- were also collected during this phase of the work. tion of all aspects of the Yanomama population is very Subsequent to this, five expediticns have been made difficult to obtain. These data must be collected from to the Yanomama tribe, in the course of which 40 vil- people whose culture places a value on recall of deceased lages we. e contacted and investigated. A few of these vil- children that differs greatly from that of our culture. In lages were contacted by non-Indians for the first time in the case of the Yanoniama, the culture actually discour- iheir history when visited by members of our group, at- ages mention of deceased children, so that information is testing to both the relative cultural purity of the tribe collected through informants other than the mother or and the remoteness of the villages. To date our informa- father. While it is possible io collect accurate data that tion includes (1) the most extensive demographic and clearly show the genealogical relationships among widely settlement pattern data available for any primitive group separated villages, it is difficult to estimate such basic at this cultural level, (2) typing of some 2,565 blood and demographic parameters as completed family size. Never- saliva specimens for 28 genetic systems, (3) some 655 theless, our data, which probably underestimate the physical examinations, (4) anthropometric measurements population, clearly indicate that the population is grow- on 402 persons, (5) dental studies on 473, and (6) der- ing. In addition, village histories and settlement data indi- matoglyphics on 488. cate that it is also expanding geographically (Chagnon, These data have resulted in a number of specific analy- 19o8a; 1968b; Neel and Chagnon, 1968). ses and publications. The high points of some of the By the time Xavante women have completed their re- initial results follow. productive performance by virtue of age or death, 33 per- Extenshre studies on serum antibodies have been percent of their children have already died prior to the ag.: formcd by the Communicable Disease Center of the U.S. of reproduction; the corresponding figure for the Yano- Public HeAth Service and by collaborators at the Vene- mama is 16 percent. These figures exclude infanticide. zuelan Institute of Scientific Investigations. in the course The higher death rate among Xavante is attributed to the of three expeditions, extensive genealogical data and fact that measles and pertuo,is have swept through them blood and saliva typings have been made on 594 Makiri- hi the past decade (SalzP.noet al.,1967), whereas at the tare Indians from 7 villages. Gene flow between these *This 16-mm film was inepared by T. Asch and N. A. Chagnon in two tribes has taken place and new genes have entered 1970. It is distributed by the National Audiovisual Center, Na- the Yanomama population at the tribal interface, often tional Archives and Records Services, Washington, D.C. 20409.

54 time the statistics were collected, the Yanornama were still "virgin" for these two diseases (Neelet al., i970). In the case of the Yanomama, the pregnancy rates im- plied by vital statistics were checked through two independent approaches: (1) abdominal palpation of as many women as possible who are estimated to be in the reproductive age, and (2) determination of urinary chorionic gonadotrophins on these same women. In a group as unacculturated as the Yanomama, each of these efforts presents a few problems. If we assume a Yanomama woman to be exposed to the risk of pregnancy from age 15 to 40, the reproductive histories suggest a child every 6 or 7 years on the average. Physical examinations of 214 Yanomarna women suggest a birth every 2.8 years, and urine studies, one every 3.8 years (Neel and Chagnon, 1968). Some of this discrepancy is almost surely due to underreporting; the remainder may be more apparent than real. The Yanomama practice abortion, at a relatively late stage of gestation, as well as infanticidethe latter immediately following delivery. An abortion or infanticide is rarely reported in a reproductive history, but these pregnancies would be detected by the supplemen- tary approaches to the question. One significant aspect of Yanomama infanticide is FIGURE 1Life-expectancy curves for a primitive cul- that more female children are killed at birth than male ture (the Yanomama), an advanced agriculturally based children. In some areas of the tribe, one third of the culture in the tropics (India in 1901-1911), and an in- adult males die in warfare. Here, a high value is placed on dustrialized culture (Japan in 1960). Deaths due to in- male children who grow up to be warriors. Accordingly, fanticide have not been included in the Yanomama curve. disproportionately more females are destroyed at birth. All three curves are for males only. Villages in these areas have a tertiary sex ratio of 1.40 compared with 1.20 for villages at the less warlike tribal reproducing at near capacity in order to maintain his periphery (Chagnon, 1968a). numbers, there is little reason to consider reproductive Figure 1 presents the Yanomama life-expectancy compensation. But with the opportunities for compen- curve. This curve represents natural causes of death only; sation that exist in the demographic pattern just de- it excludes infanticides. It is almost certain to be modi- scribed, it is obvious that the role of compensation musl fied as recently collected demographic data are incorpo- be explored. rated. For contrast, we present the curve of a tropical The findings on physical examination (Neelet al., agricultural population (India in 1901-1911, before the 1964; Weinsteinet al.,1967) indicate that, by the usual full impact of modern health measures had been felt) and stan-'ards, the Xavante and Yanomama are healthy the curve of a highly civilized, well-industrialized C01111 try people. In view of the fact that these groups kill mal- (Japan in 1960). Incidentally, there is almost surely gross formed infants at birth, the number of persons observed underreporting of infant mortality in the data from India. to have major congenital defects was surprising. Among There are two important points to be made from these a total of 287 Xavantes who were examined, there were data: 7 congenitally defective (3 severely mentally defective, Primitive populations were probably not of the so- 1 with club feet, 1 with polydactyly, and 2 with signs of called high-fertility, high-mortality type. This has great congenital heart disease). Among the first 274 Yano- significance for our thinking concerning the circum- mama examined, we have observed 7 cases of congenital stances under which the genetic polymorphisms cpme defect (6 with signs of c:mgenital heart disease, 1 achon- into being and were maintained (i.e., the mortality base droplastic dwarf). The diagnoses of congenital heart of natural selection). disease are based on auscultation. Since most children These findings indicate the possible role of repro- with serious or obvious defects die through infanticide ductive compensation in maintaining genetic polymor- or natural causes, a relatively high proportion of defec- phisms. Evidence has been developee for the occurrence tive children is indicated. of reproductive compensation in consanguineous mar- The relatively good health and nutrition of the chil- riages in Japan. As long as primitive man is thought of as dren is especially noteworthy. The high gamma globulin

55 levels of the mothers (cf. Neelet al.,1964, 1968a) imply POLYGYNY AND THE GENETIC a considerable passive transfer of immunity, and the pro- SIGNIFICANCE OF DIFFERENTIAL longed period of child nursing, about 3 years, not only FERTI LITY provides for some possible additional transfer of maternal antibodies, with a relatively smooth transition from pas- The Xavante, Yanomama, and Makiritare are polygy- sive to active immunity, but also gives the child a tine nous. In these cultures one reward for male achievement nutritional base (discussion in Neelet al.,1964; Neel and and longevity is additional wives. This appears to be the Salzano, 1967). pattern in many primitive cultures. Women appear com- In view of this picture of health among the children mitted to a pattern of child-spacing, which in Yanomama and young adults, the small number of older persons is women, living to age 40, results in a low variance in num- an enigma. The mean age in the villages of these Indians ber of reported live births. The mores of these primitive is about 20 years; only 19.2 percent of the population societies tend to minimize the variance in number of live were thought to be over 30 years of age. Studies on serum births per female but maximize the variance in number antibodies suggest early acquisition of immunity to most of children per male. of the local infectious diseases (Neel etal.,1968b); hence, It has proved quite difficult to obtain the kind of hard one would expect most of the adults to acquire immunity data that can be employed in population simulation mod- to the endemic diseases. Recent epidemics of measles and els designed to quantify the genetic consequences of pertussis among the Xavante may have killed a dispropor- polygyny. tionate number of old people, and falciparum malaria, During the past 2 years, attempts have been made to which may only now be reaching certain Yanomarna vil- develop a computer model that simulates the genetic and lages, may have had a similar effect. Nutritional disease demographic structure of the Yanomama. The basic input seems unlikely. Records of the two anthropological asso- is detailed demographic data from four villages. One of ciates contain numerous accounts of traumatic death, the objectives of this simulation is to derive an estimate through warfare or following accusations of sorcery. of the amount of inbreeding. There were great difficulties in reconciling certain aspects of the inbreeding results, after 150 simulated years, with the results of the first 20 MICRODIFFERENTIATION simulated years even though in many other respects there was a good accord with the facts. During the first 20 sim- The genetic distances between seven randomly chosen ulated years, the real population dominated the findings. Yanomama villages and seven Makiritare villages were cal- On the average, during the first 20 simulated years, the culated from data on eight gene frequencies by using the members of the real population were more closely related distance function developed by Cavalli-Sforza and to one another than were the members of the simulated Edwards. The mean distance between the Yanomama vil- population at 150 years. Even when the model specified lages is 0.330 units, and between the Makiritare villages, the maximum opportunity for consanguineous marriages 0356 units. The mean distance for the same eight gene consistent with field data, the predicted level of inbreed- frequencies between 12 Indian tribes of Central and ing declined with time. South America is 0.385 units. Thus, the average distance The reason became clear when the distribution of the between Indian villages is 85.5 to 92.4 percent of the dis- number of grandchildren per male was considered. A distance between tribes. In both of these tribes, the degree proportionate number of grandchildren are born to a few of genetic differentiation from one village to the next is males, a situation that greatly increases the possibilities felt to be noteworthy. These genetic differences are at- of full or half first-cousin marriage. Genetic significance tributed to the fact that a new Indian village usually re- is added to this phenomenon by the fact that the four sults from the fissioning of an established village. This males whose living grandchildren outnumber those of fissioning is structured by factors of kinship and tribal any male in the computer population represent two marriage practices, so that division of the village (and father-son combinations. One reason for this familial" tribal) gene pool at the time of a split is highly nonran- fertility seems to be that if, because of the polygyny of dom. Moreover, the political relationships of the groups his father, a young man possesses many sisters or half- that result from the fissioning of a village are usually sisters (who can be "traded"), that young man has an ad- hostile, and the groups tend to resettle a considerable dis- vantage in forming alliances and obtaining extra wives, so tance from each other. Thus, political forces and warfare that polygyny begets polygyny. The phenomenon does lead to a settlement movement that has a centripetal mo- not appear to be primarily genetic; rather, it reflects tion. The movement enhances the possibility that initial some of the systematic marriage and alliance formations genetic differences in recently related populations will be characteristic of tribal cultures in general and the Yano- maintained as populations migrate away from one mama in particular. One of the discoveries resulting from another. the interplay of the computer-simulation attempts and

56 -58 the demographic and cultural parameters defined by the the same at all gene frequenciesfrequency-dependent anthropologist is the surprisingly large amount of in- selection is relatively unimportant. breeding that takes place in populations such as this. Whereas a mean coefficient of relatedness of F=0.02 is generally accepted as a limit for most human populations, MEASLES actual data and simulations based on them show that this value may be greatly exceeded (MacCluer et al., in press). Immunological studies on sera collected during the The degree of differential fertility must be another 1966 expedition revealed the Yanomama to be a "virgin"' factor in the marked genetic microdifferentiation be- population with regard to measles. Measles was intro- tween villages. The number of wives a man obtains and duced into the Yanomama from Brazil just at the time of holds also depends on personal attributes, unquestionably the arrival of our 1968 expedition. There resulted a most to some extent genetically determined. Here is an exam- unusual opportunity to document (and be involved in) ple of an interaction between the genetic system and the the course of an epidemic in such a group. Observations social system. The field impression is that the polygynous suggest that the bulk of the morbidity and mortality in Indians, especially the headmen, tend to be more "intel- such populations may be ascribed to the epidemiolou of ligent" than the nonpolyunous. They also tend to have the disease under these circumstances. When all the popu- more surviving offspring. Polygyny in these tribes thus lation is ill there is a complete collapse of village life. appears to provide an effective device for certain types There is no one to gather food or bring water. Febrile in- of natural selection. dividuals dehydrate rapidly in the tropics, especially The possible genetic implications of polygyny are nursing infants whose mothers have measles. An addi- clear, but some of the facts necessary to a meaningful tional factor is the fatalistic attitude of the Indian toward treatment are still lacking. Future investigation will at- such epidemics. He tends to lie quietly doubled up in his tempt to contrast certain mental attributes of polygynous hammock awaiting death (excellent for bronchopneu- males with those of nonpolygynous males. monia).

RATE OF HUMAN EVOLUTION ESCHERICHIA COLI STUDIES

All known data on gene frequencies in various tribes Studies have been undertaken with various collabora- of American Indians were tabulated. From these data tors, at the Venezuelan Institute of Scientific Investiga- tribes were chosen for consideration on the basis of meet- tions, the U.S. Communicable Disease Center, and the ing the following criteria: (1) sample size greater than University of Michigan on the antibodies to pathogens 200, (2) non-Indian admixture estimated at less than detectable in serum specimens. Antibodies to Salmonella 5 percent, and (3) data on the six genetic loci named be- and Shigella antigens were encountered rather frequently. low plus the ABO locus. Since ail the unmixed Indians of However, diarrhea seemed a less common cause of mor- Central and South America appear to be type 0, the last- bidity and mortality than in many agricultural popula- named locus is of little value in distance measurements tions. In order to investigate this matter, in 1967, 72 but very important in opinions concerning admixture stool specimens were transported to Ann Arbor for cul- with non-Indians. ture by Dr. Warren Eveland. There were no Salmonella There were 12 tribes located in Central and South isolations. Attention thus turned to the Escherichia coli, America meeting these criteria. The genetic distances be- and 6 strains were isolated from each of these 72 speci- tween these 12 tribes, and the probable time of arrival of mens. Attempts were made to type these out with a bat- the Indian in Central and South America, have been used tery of typing sera as near complete as could be as- to estimate the maximal rate of gene substitution in these sembled. This battery would type about 99 percent of groups. The estimate was 130,000 years/gene substitution/ E. coli isolated in the United States. However, only 50 locus in any one line of descent since the arrival of the percent of these E. coli have typed out. Antisera against Indian in Central and South America. Gene substitution some of the untypable strains have been developed. At was defined in terms of the additive components of the this point at least six previously undescribed strains are vector space utilized to find the genetic distances between recognized. Within the past year, stool specimens from populations. The estimate is based on cumulative change Cali, Colombia, and Caracas, Venezuela, have been at only six genetic loci (the MNSs, Rh, Kidd, Dufy, brought back for the isolation and typing of additional Diego, and haptoglobin loci). Among the assumptions in E. coli strains. The additional samples were collected to this approach are (1) that changes in diverse directions at determine whether these new strains are widespread in various loci can be equated to directional change at one South America or characteristic of Indian populations. locus, and (2) that the coefficient of selection is about The pathogenicity of these strains is still undetermined.

59 The epidemiolou of primitive man may differ from that Goobar, W. A. Sodeman, Jr., G. H. Stollerman, E. D. Wein- of civilized man, and there is a possibility of marked local stein, and A. H. Wheeler. 1968a. Further studies of the Xavante Indians. IX. Immunologic status with respect to vari- differentiation in bacterial pressures. ous diseases and organisms. Amer. J. Trop. Med. Hyg. 17: 486-498. Neel, J. V., W. R. Centerwall, N. A. Chagnon. and H. L. Casey. 1970. Notes on the effect of measles and measles vaccine in CHROMOSOME ABNORMALITIES a virgin-soil population of South American Indians. Amer. J. Epidemiol. 91:418-429. In the course of the 1969 field work, a study of the Neel, J. V., and N. A. Chagnon. 1968. The demography of two tribes of primitive relatively unacculturated American Indians. frequency of chromosomal abnormalities was performed Proc. Nat. Acad. Sci. U.S.A. 59:680-689. in three Yanomama villages and one Piaroa village. The Neel, J. V., W. M. Mikkelsen, D. L. Rucknagel, E. D. Weinstein, objective was to obtain a base line for chromosomal dam- R. A. Goyer, and S. H. Abadie. 1968b. Further studies of the age in a population essentially unexposed to radiation, Xavante Indians. VIII. Some observations on blood, urine. and stool specimens. Amer. J. Trop. Med. Hyg. 17:474-485. pesticides, and the like. Heparinized blood samples ob- Neel, J. V., and F. M. Salzano. 1967. Further studies in the tained in villages near airstrips were refrigerated and Xavante Indians. X. Some hypotheses-generalizations resulting flown to a temporary cytogenetics laboratory established from these studies. Amer. J. Hum. Genet. 19:554-574. Neel, J. V., F. M. Salzano, P. C. Junqueria, F. Keiter; and D. at the Venezuelan Institute for Scientific Investigation. Maybury-Lewis. 1964. Studies on the Xavante Indians of the The samples usually arrived within 36 hours of collection. Braxilian Mato Grosso. Amer. J. Hum. Genet. 16:52-140 They were cultured, and preparations were made that Salzano, F. M., J. V. Neel, and D. Maybury-Lewis. 1967. Further studies on the Xavante Indians. I. Demographic data on two were read in Ann Arbor. Where possible, 100 cells were additional villages; genetic structure of the tribe. Amer. J. scored per individual. Members of the expedition served Hum. Genet. 19:463-489. as controls. An effort was made to assemble an age- and Weinstein, E. D., J. V. Neel, and F. M. Salzano. 1967. Further studies on the Xavante Indians. VI. Physical status of the sex-stratified sample of largely unrelated individuals. The Xavantes of Simoes Lopes. Amer. J. Hum. Genet. 19:532-542. amount of damage is clearly greater than was expected. Weitkamp, L. R., and N. A. Chagnon. 1968. Albumin Miku. Some cells show striking damage. Present data indicate A new variant of human serum albumin. Nature 217:759-60. that highly complex chromosomal aberrations are found in individuals of all ages and of both sexes. In the future we will extend our collecting of genetic and anthropological data to Yanomama villages that have APPENDIX A: PUBLICATIONS not been studied. We will initiate auxiliary studies on the Yanomama, Arends, T. G., G. Brewer, N. Chagnon, M. Gallango, H. Gershowitz, M. Layrisse, J. Neel, D. Shreffler, R. Tashian, including ethnobotanical studies and dental studies. An and L. Weitkamp. 1967. Intratribal genetic differentiation ethnobotanist and a zoologist will be added to the field among the Yanomama Indians of Southern Venezuela. Proc. team. The ethnobotanist will explore the botanical re- Nat. Acad. Sci. U.S.A. 57:1252-1259. Chagnon, N. A. 1970. The culture-ecology of shifting (pioneering) sources of the tropical forest culturally significant to the cultivation among the Yanomama Indians, p. 249-255. In Yanomama. The zoologist will do the same for the pro- Proceedings of the VIIIth International Congress of Anthro- tein aspects of Yanomarna economic productivity. pological and Ethnological Sciences. Vol. 3. Tokyo. Chagnon, N. A., P.-LeQuesne, and I. Cook. 1971. Yanomama We intend to initiate the same kind of multidisciplin- hallucinogens: anthropological, bacterial and chemical find- ary and multinational studies among several other South ings. Cum Anthropol. 12:72-74. American tribes. One of these tribes was contacted for Fitch, W. M., and J. V. NeeL 1969. The phylogenetic relationships of some Indian tribes of Central and South America. the first time late in 1969 and is totally unknown. Amer. J. Hum. Genet. 21:384-397. Neel, J. V. 1969. Some aspects of differential fertility in two American Indian tribes, p. 356-361.In Proceedings of the VIIIth International Congress of Anthropological and Ethno- logical Sciences. Vol. 1. Tokyo. REFERENCES Neel, J. V. 1969. Some changing constraints on the human evolutionary process, p. 389-403. In Proc. XIIth Int. Congr. Chagnon, N. A. 1968a. Yanomama: the fierce people. Holt, Genet. Vol. 3. Rinehart, and Winston, New York. Neel, J. V., and R. H. Ward. 1970. Village and tribal genetic dis- Chagnon, N. A. 1968b. Yanomama social organization and war- tances among American Indians, and the possible implications fare, p. 109-159. In M. Fried, M. Harris, and R. Murphy for human evolution. Proc. Nat. Acad. Sci. U.S.A. 65:323-330. [ed.] War: the anthropology of armed conflict and aggession. Post, R. H., J. V. Neel, and W. J. Schull. 1968. Tabulations of Natural History Press. phenotype and gene frequencies for 11 different genetic sys- Changon, N. A., J. V. Neel, L. Weitkamp, H. Gershowitz, and tems studied in the American Indian, p. 141-185. In Bio- M. Ayres. 1970. The influence of cultural factors on the medical challenges presented by the American Indian. Sci. demography and pattern of gene flow from Makiritare to the Pub. 165. Pan American Health Organization, Washington, Yanomama Indians. Amer. J. Phys. Anthropol. 32:339-47. D.C. MacCluer, J. W., J. V. Neel, and N. A. Chagnon. In press. Demo- Schull, W. J., and J. W. MacCluer, 1968. Human genetics: struc- graphic structure of a primitive population: a simulation. ture of populations. Annu. Rev. Genet. 2:279-304. Amer. J. Phys. Anthropol. Ward, R. H., and J. V. NeeL 1970. Gene frequencies and micro- Neel, J. V., A. H. P. Andrade, G. E. Brown, W. C. Eveland, J. differentiation among the Makiritare Indians. IV. A compari-

58 son of a generic network with ethnohistory and migration Henry Gershowitz, Department of Human Genetics,Univers4 matrices; a new index of genetic isolation. Amer. J. Hum. of Michigan Medical School, Ann Arbor (geneticist) ' Genet. 22:538-561. Ronald Griffith, Department of Human Genetics, University of Michigan Medical School, Ann Arbor (computer programmer) Miguel Layrisse, Instituto Venezolano de Investigaciones Cien- tificas, Apartado 1827, Caracas, Venezuela (geneticist) Philip LeQuesne, Department of Chemistry, University of Michi- gan, Ann Arbor (chemist) Jean MacCluer, Department of Human Genetics, University of APPENDIX B: PARTICIPANTS Michigan Medical School, Ann Arbor (ma thematician- James V. Neel, Department of Human Genetics, University of geneticist) Michigan Medical School, Ann Arbor (Program Director; D. Maybury-Lewis, Deparfment of Social Relations, Harvard geneticist) University, Cambridge, Mass. (cultural anthropologist) Timothy Asch, 4 Bryant Street, Cambridge, Mass. (ethno- Ernest Migliazza, Caixa Postal 165, Boa Vista, Territorio Federal graphic film maker) de Roraima, Brazil (linguist) Manuel Ayres. Travessa Presidente Pernambuco, 104, Belem, William J. Oliver, Pediatrics Department, Mott Hospital, Ann Para, Brazii (serologist) Arbor, Mich. (pediatrician-geneticist) H. Powell Baker, 132 School Street, Manchester, Mass. (pedia- Francisco M. Salzano, Departamento de Genetica, Universidade trician-geneticist) de Rio Grande do Sul, Caixa Postal 1953, Porto Alegre, Rio G. H. Bergold, Chief, Virology Department, Instituto Venezo- Grande do Sul, Brazil (geneticist) lano de Investigaciones Cientificas, Apartado 1827, Caracas, William J. Schull, Department of Human Genetics, University Venezuela (virologist) of Michigan Medical School, Ann Arbor (mathematician- Arthur Bloom, Department of Human Genetics, University of geneticist) Michigan Medical School, Ann Arbor (cytogeneticist) Donald C. Shreffler, Department of Human Genetics, University Helen Casey, Chief, Viral Immunoserology Unit, National Com- of Michigan Medical School, Ann Arbor (geneticist) municable Disease Center, U.S. Department of Health, Edu- Richard E. Tashian, Department of Human Genetics, University cation, and Welfare, Atlanta, Ga. (epidemiologist) of Michigan Medical School, Ann Arbor (geneticist) Willard Centerwall, Department of Pediatrics, Loma Linda Uni- R. H. Ward, Department of Anthropology and Medical School, versity School of Medicine, Loma Linda, Calif. (pediatrician- University of Auckland, Private Bag, Auckland, New Zealand geneticist) (geneticist) N. A. Chagnon, Department of Human Genetics, University of E. David Weinstein, 4630 Browndeer Lane, Palos Verdes Penin- Michigan Medical School, Ann Arbor (cultural anthropolo- sula, Calif. (pediatrician) gist) Lowell R. Weitkamp, Division of Genetics, University of Warren C. Eve land, School of Public Health, University of Rochester School of Medicine, Rochester, N.Y. (pediatrician- Michigan, Ann Arbor (epidemiologist) geneticist)

59 61 Biology of Human Populations at High Altitudes

Exploring the stresses associated with high altitude is investigators from several disciplines in the United States one of the best approaches to determining the limits of and Peru. Although the major proposals have not been human ability to adapt to environmental stresses. A good funded, smaller research projects on related topics have experimental situation is provided by the fact that at progressed without interruption. high altitude the stress level produced by hypoxia is extreme. Technological change has not substantially ame- liorated the effect of the lower atmospheric pressure. This contrasts with stress such as that produced by cli- OBJECTIVES matic extremes in temperature where technology can reduce or even eliminate the stress. Also, the hypox.:as related to altitude appear to have significant effects on the manifestations of several degenerative diseases that are of The general objective of the program is to obtain an major importance to a large segment of the world popu- understanding of how human populations at high altitude lation. adapt to their environment and of what happens to their Human survival, health, and productivity all appear to biology when they migrate downward. Since this general be modified significantly by residence at high altitude. objective is too broad for complete exploration during The apparent reduction in fertility at high altitude is the tenure of the IBP, certain aspects are being empha- noteworthy in view of the current population explosion sized. elsewhere. Among populations living in the Andes, blood How does residence in the high altitude environ- pressure is lower and fatal heart attacks appear less fre- ment modify the incidence, severity, and natural history quent than in populations at sea level. This suggests that of human degenerative cardiovascular diseases (e.g., sys- the high altitude environment may afford some protec- temic hypertension, myocardial infarction, pulmonary tion from degenerative cardiovascular diseases, which are thromboembolism, and stroke)? the major causes of death in the United States today. What is the incidence of maladaptation to high alti- However, a man migrating to high altitude must ac,ept tude, (e.g., excessive polycythemia, chronic mountain the fact that his work output will be less than if he stays sickness, acute pulmonary edema, central nervous system at sea level. These are but a few of the observations that dysfunction)? How frequently is intolerance to altitude have prompted this study of man's adaptability to the the major factor in the decision to leave the high altitude environmental stress of high altitude. environment? Co-directors of th::, program are Paul T. Baker, Penn- How is reproductive capacity reduced in human sylvania State University; Robert F. Grover, University populatons at high altitude? Are there fewer pregnancies of Colorado; and John Rankin, University of Wisconsin. (male or female infertility), more spontaneous abortions, Dr. Bake: began studying the Indian populations of the fewer live births? Are there any differences in the inci- high Andes in 1962 and established a permanent field dence of congenital malformation or infant mortality? station in Nufioa, Peru, at 4,000 m, which has functioned Does altitude modify human growth and develop- continuously since 1964. Dr. Grover began his studies of ment? the population of Leadville, Colorado, at 3,100 m, in the Do native high altitude populations show evidence Rocky Mountains in 1961 and established a permanent of better biological adaptation to altitude than upward laboratory there in 1964. Together with Dr. Rankin, they migants, and if so at what stage are these adaptations submitted the pilot description of this program in the manifest? spring of 1968 and submitted major proposals later that Do downward migrants develop health problems year. The proposals were a logical extension of well- that are specifically related to their previous adaptation founded research programs involving the collaboration of to altitude?

62 RESEARCH PLANS standards are similar to those in many low-altitude populations in the United States, nonaltitude factors are minimized. This fact simplifies the task of identifying the ef- For several reasons the scientific goals of this program fects of altitude per se. The entire study is made feasible can be achieved only by conducting several carefully co- by community acceptance and cooperation. ordinated subprograms. Data from ongoing research will be used in the sub- "High altitude" is not a single elevation above sea program dealing with Framingham and Tecumseh. level, but a continuum. Altitude as a significant physio- A comparison of two communities at the same alti- logical stress begins at about 10,000 ft and extends to tude, one in the United States and one in the Andes (sub- about 17,000 ft, the upper limit tolerable for permanent programs 1 and4),will serve to evaluate genetic factors human residence. operating through many generations. Human adaptation to high altitude occurs in varying Subprograms 3 and 5 will provide maximum contrast degrees and is often less than complete. Factors involved between the effects of hypoxia on genetically similar include altitude itself; a person may adapt completely to populations, 2:nd they will show the effects of downward 10,000 ft but not to14,000ft. Age is important; young migration of a population with maximal high-altitude people adapt more readily than adults. Women appear to acclimatization. have a better physiological adaptation than men. There Subprograms 1,4,and 5 will provide maximum con- is always a spectrum of physiological responses among trast between the effects of factors other than hypoxia individuals; this is greatest in people from sea level who (e.g., diet, race, and physical activity) that may be im- move to altitude. portant in explairing the lower incidence of cardiovascu- Man's tolerance of a particular high altitude environ- lar disease in the Andes. ment depends not only on physiological response to hy- In each community, a large amount of biomedical and poxia but also on a variety of biological and cultural fac- social data will be collected on each member of the popu- tors. Differences in the diet, social expectations, lation sample. The data will be collected in a form that child-rearing practices, and disease treatment are a few will be suitable for computer processing. This will permit factors that may affect responses to altitude. The factors efficient analysis, cross comparisons, and ultimate can be evaluated only by comparing them in several synthesis. communities. Because of these factors, interpopulation and intra- population comparisons are needed. Data for the comparisons will be gathered in five subprograms: RESEARCH METHODS AND PROGRESS 1. Study of the population of Leadville, Colorado, in the Rocky Mountains. The altitude is 10,000 ft (moderate). 2. Study of two sea-level communities in the United Extensive groundwork has been laid for the proposed States: Framingham, Massachusetts, and Tecumseh, Mich- studies; persons and organizations skilled in population igan. These communities have been under investigation surveys have been contacted at the University of Wiscon- for several years, and considerable information is avail- sin. Detailed plans for a special census of Leadville have able. been completed. Only funding is needed to activate these 3. Study of a sea-level community in the Andes plans. Cocrachra, Peru. The base of operations in Leadville is the High Alti- 4. Study of a community at moderate altitude in the tude Research Laboratory in St. Vincent's Hospital. This Andes (not yet selected). laboratory was established in1964by Dr. Grover and has 5. Study of a community at high altitude in the been funded by the National Institutes of Health (NIH) AndesNufioa, Peru (13,000 ft). for the past 7 years. Continued support has been re- The subprogram dealing with the population of Lead- quested. ville, Colorado, will be comprehensive and sophisticated. In Nuiloa, Peru, community research is conducted Knowledge about the capacity for long-term adaptation from a research laboratory that Dr. Baker constructed to altitude among persons of European ancestry is of there in1964.Activities are coordinated in Nufioa and great importance, since most available data were obtained in Lima by Tulio Velasquez, San Marcos University. on Andean Indians. The desired knowledge can be gained Despite the lack of comprehensive funding, there has at Leadville. The oldest families have been there for four been progress in some aspects of the research. The ap- generations. One finds not only natives representing these pended list of publications indicates the general nature of families but also newcomers of various ages, some of results to date. The following paragraphs refer to specific whom have lived at altitude for many years. Since living results achieved during1969.

61 Working Capacity reduces the oxygen pressure in the blood. When the oxygen pressure is already lowered at high altitude, the The physical working capacity of a group of Leadville further reduction caused by smoking becomes very sig- natives and comparable groups of sea-level natives was de- nificant. At high altitude (1,:-adville), an excessive thick- termined at both high and low aititudes. The working ening of the blood (polycythemia) sometimes results. If capacity of the two groups was identical at sea level and the affected persons stop smoking, the polycythemia dis- was reduced equally (25 percent) in both groups at alti- appears in 3 months. Thus, in some persons, an apparent tude. Thus, the Leadville native exhibits no advantage in maladaptation to high altitude (polycythemia) may be capacity for physical work at altitude in spite of accli- caused by cigarette smoking. matization to altitude from birth. (This conclusion is in Participants: George J. Brewer, John Eaton, John V. contrast with the apparent advantage enjoyed by the Weil, and Robert F. Grover. Andean native.) Participants: John T. Reeves and Robert F. Grover. Fertility and Neonatal Mortality Diffusing Capacity of the Lung Demographic data from a high-altitude Peruvian popu- The capacity of the lung to transfer oxygen from air lation suggest that extreme altitude may have a slightly to blood (pulmonary diffusing capacity) was measured depressant effect on fertility and certainly affects neo- in men and women at sea level, in natives of Leadville, natal mortality. Analysis on the project is continuing. and in newcomers adapted to high altitude. Diffusing Participants: Paul T. Baker and James S. Dutt. capacity at high altitude was sipificantly greater in the natives of Leadville but not in the newcomers, implying Effects on Growth that lung structure is augmented by development at high The study of the effects of altitude on the growth altitude. of high-altitude natives in Peru suggests that the rates of Participants: Robert L. Johnson, Arthur C. De Graff, growth may be slowed at extreme altitudes. However, John M. Miller, and Robert F. Grover. the study of downward Indian migrants indicates that the unusually large chest characteristic of high-altitude Performance of the Heart natives may be genetically fixed. Second-generation na- The quantity of blood pumped by the heart per beat tives at low altitude had adult chest measurements com- (stroke volume) and per minute (cardiac output) was parable with those of genetically similar natives at alti- measured in Leadville residents and in comparable sea- tude. Study of this problem, including the relationship level residents at both high and low altitudes. Both stroke between chest measurements and lung function, is con- volume and cardiac output are reduced at high altitude tinuing. both in newcomers and in adapted long-term residents. Participants: A. Roberto Frisancho, Charles J. Hoff, This is probably the major factor causing the reduction and Paul T. Baker. in working capacity at altitude. (These fmdings are in contrast with data from Andean natives, whose cardiac Responses to Cold output at altitude is not reduced.) A series of studies of responses to cold have been Participants: Loren H. Hartley, James K. Alexander, completed. These studies indicate that the high-altitude and Robert F. Grover. native has unusually high peripheral blood flow in the presence of exposure to cold. Coca chewing does not ap- Coronary Circulation pear to affect temperature regulation as much as it does Because cardiac output is reduced at high altitude, psychophysiological responses to exercise. while blood pressure remains unchanged, the work of the Participants: Michael A. Little and Joel M. Hanna. heart is reduced. Thus, the heart muscle requires less oxygen, and the demands on the coronary circulation are less (coronary blood flow is reduced at high altitude). These APPENDIX A: PUBLICATIONS fmdings have potentially beneficial implications for per- Baker, P. T. 1967. The study of biological adaptive mechanisms sons with coronary artery disease. in a high altitude population. Amer. J. Phys. Anthropol. Participants: James K. Alexander, Roberto Lufsha- 27:242. (Abstr.) rowski, and Robert F. Grover. Baker, P. T. 1969. Human adaptation to high altitude. Science 163:1149-1156. Baker, P. T., G. Escobar, G. DeJong, C. J. Hoff, R. B. Mazess, Effect of Cigarette Smoking J. Hanna, M. A. Little, and E. Picon-Reatigui. 1968. High altitude adaptation in a Peruvian community. Occasional Cigarette smoking not only reduces the capacity of papers in anthropology No. 1. Department of Anthropology, the blood to carry oxygen (carboxy hemoglobin); it also The Pennsylvania State University.

62 Baker. T. S., A. V. Little. and A. R. Frisancho. 1967. Infant Little. M. A. 1967. Foot temperature responses to a moderate growth in a high altitude population. Amer. J. P]:vs. Anthro- cold stress in a sample of highlr.nd Peruvian Indian males. pol. 27:248. (Abstr.) Amer. J. Phys. Anthropol. 27:248. (Abstr.) Brewer, G. J.. J. Eaton. J. V. Weil, and R. F. Grover. 1970. Mazess, R. B. 1967. Exercise performance at altitude. Amer. J. Studies of red cell glycolysis and interactions with carbon Phys. Anthropol. 27:247. (Abstr.) monoxide, smoking and altitudes. p. 95-114. /n G. J. Brewer Mazess, R. B. 1968. Hot-cold food beliefs among Andean peas- [ed. I Advances in experimental medicine and biology. Vol. 6. ants. J. Amer. Diet. Ass. 53(2):109-113. Plenum, New York. Mazess, R. B. 1968. The oxygen cost of breathing in man: effects Buskirk, E. R. 1969. Decrease in physical working capacity at of altitude. training and race. Amer. J. Phys. An thropol. high altitude. In Biomedicine of high terrestrial elevations. 29(3):365-376. U.S. Army Research Institute of Environmental Medicine. Mazess, R. B. 1969. Altitude and adaptation. Amer. J. Phys. Natick, Mass. A nthropol. 31(2):259. (Abstr.) Buskirk. E. R., J. Kollias, R. F. Akers, E. K. Prokop, and E. Mazess, R. B. 1969. Enercise performance at high altitude in Picon-Reatigui.967. Maximal performance at altitude and Peru. Fed. Proc. 28(3):1301-1306. on return from altitud: in conditioned runners. J. Appl. Mazess, R. B. 1969. Exercise performance of Indian and white Physiol. 23:259-266. high altitude residents. Hum. Biol. 41(4):494-518. Buskirk. E. R., and J. Men lez. 1967. Nutrition, environment and Mazess, R. B. 1970. Cardiorespiratory characteristics and adap- work performance, with special reference to altitude. Fed. tation to high altitudes. Amer. J. Phys. Anthropol. 32(2): Proc. 26:1760-1767. 267-278. DeGraff, A. C.. Jr.. R. F. Grover, R. L. Johnson, Jr., J. W. Mazess, R. B., E. Picon-Reatigui, R. B. Thomas, and M. A. Hammond, Jr., and J. M. Miller. 1970. Increased diffusing Little. 1968. Effects of alcohol and altitude on man during capacity of the lung in persons native to 3,100 m in North rest and work. Aerosp. Med. 39:403-406. America. J. AppL Physiol. 29:71-76. Mazess, R. B., E. Picon-Reatigui, R. B. Thomas, and M. A. Dun. J. S. 1970. The effect of cultural and geo6raphic factors Little. 1969. Oxygen intake and body temperature of basal on human movement ..nd gene flow in a highland Peruvian and sleeping Andean natives at high altitude. Acrosp. Med. Quechua community. Amer. J. Phys. Anthropol. 33:(1):128. 40(1):6-9. (Abstr.) McClung, J. P. 1969. The effects of high altitude on human Frisancho, A. R. 1969. Morphological variation during growth birth. Harvard Univ. Press, Cambridge, Mass. related to altitude hypoxia. Amer. J. Phys. Anthropol. Thomas, R. B. 1970. Body size and energy flow in a high 31(2):260. (Abstr.) Andean population. Amer. J. Phys. AnthropoL 33(1):144. Frisancho, A. R. 1970. Developmental responses to high altitude (Abstr.) hypoxia. Amer. J. Phys. Anthropol. 32(3):401-408. Thomas, R. B. 1967. Hand temperature responses of Peruvian Frisancho, A. R., and P. T. Baker. 1970. Altitude and growth: Indians to localized cold exposure. Amer. J. Phys. Anthropol. a study of the patterns of physical growth of a high altitude 27:248. (Abstr.) Peruvian Quechua population. Amer. J. Phys. Anthropol. Weitz, C. A. 1970. Morphology and total body cooling among 32(2):279-292. Quechua Indians. Amer. J. Phys. Anthropol. 33(1):146. Frisancho, A. R., M. T. Newman, and P. T. Baker. 1970. Dif- (Abstr.) ferences in stature and cortical thickr.ess among highland Quechua Indian boys. Amer. J. Clin. Nutr. 23(4):382-385. Gai:uto, R. M. 1970. Pulmonary function and body morphology: selected relationships studied at high altitude. Amer. J. Phys. Anthropol. 33(1):130. (Abstr.) APPENDIX B: PARTICIPANTS Grover, R. F. 1968. The high altitude resident of North America. Scientia (Milan) 103:1-17. Paul T. Baker, Department of Anthropology, The Pennsylvania Gursky, M. J. 1970. Diet and physical characteristics of Quechua State University, University Park (Co-Director of Program) Indians from three Peruvian highland communities. Amer. J. Robert F. Grover, Research Director, Cardiovascular Pulmonary Phys. Anthropoi. 33(I):131. (Abstr.) Research Laboratory, University of Colorado Medical Center, Hanna, J. M. 1969. A comparison of laboratory and field studies Denver (Co-Director of Program) in detecting variation in response to cold. Amer. J. Phys. John Rankin, Rennebohm Professor of Medicine, University of Anthropol. 31(2):258. (Abstr.) Wisconsin Medical Center, Madison (Co-Director of Program) Hanna, J. M. 1970. A comparison of laboratory and field studies James K. Alexander, Department of Cardiology, Baylor Univer- of cold response. Amer. J. Phys. AnthropoL 32(2):227-232. sity College of Medicine, Texas Medical Center, Houston Hanna, J. M. 1970. The effects of coca chewing on exercise in George J. Brewer, Department of Human Genetics, University of the Quechua of Peru, Hum. Biol. 42(1):1-11. Michigan, Ann Arbor Hartley, L. H., J. K. Alexander, M. Modelski, and R. F. Grover. Elsworth R. Buskirk, Human Performance Laboratory, The 1967. Subnormal cardiac output at rest and during exercise in Pennsylvania State University, Univergity Park residents at 3,100 m altitude. J. Appl. Physiol. 23:839-848. Arthur C. De Graff, Laboratory of Cardio-Pulmonary Research, Hultgren, H. N., and R. F. Grover. 1968. Circulatory adaptations Department of Internal Medicine, Southwestern Medical to high altitude. Annu. Rev. Med. 19:119-152. School, University of Texas, Dallas Kollias, J., E. R. Buskirk, R. F. Akers, E. K. Prokop, P. T. Gordon F. DeJong, Department of Sociology, The Pennsylvania Baker, and E. Picon-Reatigui. 1968. Work capacity of long- State University, University Park time residents and newcomers to altitude. J. Appl. Physiol. James S. Dutt, Department of Anthropology, The Pennsylvania 24:792-799. State University, University Park Little, M. A. 1969. Effects of alcohol and coca on foot tem- John Eaton, Department of Human Genetics, University of perature responses of highland Peruvian Indians during a Michigan, Ann Arbor localized cold exposure. Amer. J. Phys. Anthropol. 31(2):259. Gabriel Escobar, Department of Anthropology, The Pennsyl- (Abstr.) vania State University, University Park Little, M. A. 1969. Temperature regulation at high altitude: A. Roberto Frisancho, Center for Human Growth and Develop- Quechua Indians and U.S. whites during foot exposure to ment, University of Michigan, Ann Arbor cold water and cold air. Hum. Biol, 41(4):519-535. Ralph M. Garruto, Department of Anthropology, fhe Pennsyl- Little, M. A. ; 970. Effects of alcohol and coca on foot tem- vania State University, University Park peratures of highland Peruvians during a localized cold ex- Martin J. Gursky, Department of Anthropology, The Pennsyl- posure. Amer. J. Phys. AnthropoL 32(2):233-242. vania State University, University Park

65 Joel M. Hanna, Department of Physiology. University of Hawaii, Jean P. McClung, Harvard University Medical School, Cambridge, Honolulu Mass. Capt. Loren H. Hartley, U.S. Army Research Institute of Envi- John M. Miller, Laboratory of Cardio-Pulmonary Research, ronmental Medicine, Natick, Mass. Department of Internal Medicine, Southwestern Medical Charles J. Hoff, Department of Anthropology. University of School, University of Texas, Dallas Oregon, Eugene Emilio Picon-Reatigui, Institute of Andean Biology, San Marcos Robert L. Johnson, Cardio-Pulmonary Laboratory, Department University, Lima, Peru of Medicine, Southwestern Medical School, University of John T. Reeves, Director, Cardiovascular Laboratory, University Texas, Dallas of Kentucky Medical Center, Lexington James Kollias, Human Performance Laboratory, The Pennsyl- R. Brooke Thomas, Department of Anthropology, Cornell Uni- vania State University, University Park versity, Ithaca Michael A. Little, Department of Anthropology, The Ohio State Tulio Velasquez, Institute of Andean Biol NY. San Marcos Uni- University, Columbus versity, Lima. Peru Roberto Lufsharowski, Department of Cardiology, Baylor Uni- John V. Weil, Assistant Professor of Medicine and Assistant versity College of Medicine, Texas Medical Center. Houston Director, Cardiovascular-Pulmonary Research Laboratory, Richard B. Mazess, Department of Anthropology. University of University of Colorado Medical Center, Denver Wisconsin, Madison Charles A. Weitz, Department of Anthropology, The Pennsyl- Jose Mendez, Human Performance Laboratory, The Pennsylvania vania State University. University Park State University, University Park

64 International Study of Circumpolar Peoples

Eskimos, similar in race, lancruage, and culture across RESEARCH PLANS a distance of over 4,000 miles at the top of the world, represent one of the greatest linear dispersions of closely rekited peoples in the history of mankind. Their habitat, Research on Eskimos has been in five categories. The F.low-covered for two thirds of the year, is characterized categories and the main components of each are as fol- by low temperatures, seasonal extremes in light and dark- lows: ness, and relatively meager sustaining resources. Despite General health and physical fitness Cardiovascular these environmental constraints, Eskimos have occupied disease, infant morbidity and mortality, and epidemiology the Arctic for probably 400 generations. Their long-term Growth Dentition, size and shape of the body at dif- occupancy in small, kin-based hunting groups provides ferent ages, and the mineral content of bone human biologists with unique opportunities to study in Genetics Population genetics and demography detail certain aspects of human adaptability to a harsh Behavior Cognition, psychomotor and expressive be- environment. Prior to c. 7,000 B.C., most of our own havior, and behavioral genetics evolutionary development occurred under similar circum- Ecology Nutrition and studies of energy flow through sonces in a more southerly setting. The Eskimo way of the ecosystem of which man is a part life is changing radically and rapidly in response to social Three years of field research have been conducted. and economic pressures that are growing with develop- Nineteen scientists started work at Wainwright, Alaska, ment of mineral and other natural resources in the Es- in July 1968. They were from four American universities, kimos' home territories. Therefore, it is imperative that the Alaska State Department of Health, the U.S. Public biological base-line information be obtained before it is Health Service, and the University of Milano, Italy. too late. During the summer of 1968, 91 percent of the Eski- mos in Wainwright were examined. Medical examinations included electrocardiographs; X rays of chest, head, wrists, and ankles; tests of the n-dneral content of bone; OBJECTIVE blood tests; and throat cultures. Dental examinations included X rays, dental casts, and saliva collections. Respi- ratory parameters, endurance, and body rhythms were The objective of this program is to describe biological measured. In addition, anthropolocal materials were and behavioral processes responsible for the successful collected; they included genealogical data, family histo- adaptation and perpetuation of breeding isolates of Es- ries, and data obtained by photography, anthroposcopy, kimos in northern Alaska. and anthropometry. Villages in northern Alaska were chosen for this study Most investigators used equipment and supplies from because they have an optimum size for study, the vil- their own research laboratories, and subsequent analyses lagers have retained hunting and fishing as a way of life, were supported by the home institutions. The Alaska and information from previous investigations is accessible. State Department of Health provided X-ray equipment. The Alaskan studies are coordinated with IBP studies A nutdtional survey was made in January and Febru- being conducted in other parts of the Arctic by Danish, ary 1969. The investigators estimated amounts of dif- French, and Canadian investigators. Methods are com- ferent types of food that were eaten, measured hemo- parable, and results can readily be synthesized. globin and hematocrit values, and collected (for later laboratory assay) specimens of urine from children of school and preschool age, selected families, and members The Director of the program on circumpolar peoples has been on an extended absence. Only the part of the program that is con- of the National Guard. cerned with Eskimos is described in this report. Work was continued in the summer of 1969 with

65 support from the National Institute of General Medical Minnesota for analysis and will provide a good base line Sciences. Those participating included dentists, chrono- for the present study. biologists, psychologists, epidemiologists, exercise physiologists, an anthropometrist, a dermataglyphic specialist, a physician, an ophthalmologist, a bone physicist, and a SERUM EPIDEMIOLOGY demographer. In addition to carrying out his research, the ophthalmologist, who was from Finland, refracted Two hundred specimens of whole blood (30 ml each) eyes and prescribed glasses for 50 persons. The Chief of were obtained by venipuncture from all persons between Ophthalmology at the Alaskan Native Health Center in the ages of 6 and 74. All blood samples were slide-typed Anchorage provided extra equipment in exchange for for blood types A, B, 0 and MN to aid in later identifica- these services. tion. The specimens of whole blood were centrifuged, Additional work was done at Anaktuvuk Pass (in the and the serum was removed asceptically. The serum was Brooks range), where Wainwrighters have relatives. The distributed in 0.5-ml aliquots and stored in liquid nitro- anthropologists obtained anthropometric measurements gen. About 2,100 vials of packed red cells and 1,000 vials and finger and hand prints of 23 women and 13 men at of plasma were processed and stored in the summer of the village, thus increasing the sample size. 1968, and 1,000 vials of plasma were processed and During the summer of 1969 a psychiatrist undertook stored in 1969. the first of a series of regular visits to Wainwright, and Blood specimens were shipped to the New York City the chronobiologist continued his work during September. Blood Bank for detailed blood type, genetic marker, and Frederick A. Milan is Director of the subprogram con- enzyme studies. Serum samples (180) were sent to the cerned with the study of Eskimos. Case Western Reserve University for study of serum amylase isoenzymes in relation to salivary amylase isoenzymes and typing of immunoglobins. As part of another study, RESEARCH PROGRESS serum samples (185) are being analyzed for C 3 "complement" component at the Department of Medicine, University of Wisconsin. Serum samples (190) were ana- The following statements about the Eskimo popula- lyzed by the Clinical Chemistry Section, Wisconsin State tion at Wainwright are based on the work performed in Laboratory of Hygiene, for standard blood chemistry 1968 and 1969. profiles. Sera (190) are being studied for arborvirus antibodies. Other studies are being undertaken for antibodies of pertussis, mononucleosis, measles, mumps, rubella, GENERAL HEALTH tularemia, brucellosis (especially rangifer0 and mycotic agents. Comparisons will be drawn between these data In general, the population is healthy, but the incidence and data obtained from a comparable study of a native of chronic otitis media is high (45 percent of adults, 40 group at Xuixquilucan, state of Mexico, Mexico. percent of children), and there are marked seasonal fluctuations in upper respiratory infections. Infection from past exposure to puhnonary tuberculosis (45 percent of WORK CAPACITY population) is now controlled chemoprophylactically. Only six persons have chronic diseases; yet infant mor- The physical fitness of 62 Eskimo males (age 6 to 64 bidity and mortality rates are extremely high_ Serum years) was evaluated by means of a submaximal step test, cholesteral levels (mg/229A ml) are in agreement with by a test to measure maximal anaerobic power, and by a levels from the United States. 15-min endurance run. Certain clinical respiratory tests (vital capacity, timed vital capacity, and maximal breathing capacity) were also undertaken. CARDIOLOGY The results indicate that peak maximal aerobic power of Eskimo males occurs between the ages of 15 and 20 Electrocardiograms were sent to the Univer3ity of and is similar in magnitude to that of other population Minnesota for standard analysis and computer-compatible groups. The decline of maximal aerobic power with age coding. Serum cholesterol and triglyceride levels are avail- also is similar to that of other population groups. able and will be correlated with incidence of heart disease. Endurance can be measured in a run lasting 15 min. The incidence of atherosclerotic heart disease in natives The only instructions are to run as far as possible. Un- of Alaska is one eighth that in our population. Electro- fortunately, the Eskimos sprinted at the start and ex- cardiograms were obtained on this same population in hausted themselves, and their performance could not be 1955. These data have also been sent to the University of evaluated by this method.

66 Vital capacity and maximal breathing capacity are posterior (Arcus senilis), the cilioretinal artery, the pig- 10 percent higher in Eskimo males than capacities pre- ment content of the conjunctiva and eye gound, the dicted from height and weight standards for Eskimos in form of the palpebral fissure, and the eyelid fold. He took general. The relatively high value of seated height in the color pictures of the irises and fundi of the eyes of 98 ratio between seated height and stature could account persons. for some of this elevation. In this population there were three blind persons, one In summary, the Eskimo male population is similar to of whom had narrow angle glaucoma. There were two other populations in their capacity to perform aerobic cases of squint with amblyopia and two casesof color work but above average in their capacity for short bursts blindness. of muscular power. It would appear that centuries of Anatomically, the anterior chamber of the eye was adaptation to an arctic hunting life has not led to devel- shallower than in Europeans but not as shallow as in Es- opment of a significantly more efficient oxygen transfer kimos in the Upernavik district of Greenland. A shallow system by these people. Nevertheless, 80 percent of this anterior chamber predisposes elderly Eskimos to narrow Eskimo sample are physically fit as compared with a angle glaucoma. The climatological changes of the con- comparable U.S. sample, in which only 20 percent were junctiva, such as development of the pinguecula and fit. changes of the corneal epithelium (band keratitis), were also less pronounced than in the Eskimos L-1 Greenland or the Lapps in Finland. CHRONOBIOLOGY About 50 of the 308 persons living in the village received prescriptions for new eyeglasses as a side benefit The periodicity of light intensity in the polar region is for their cooperation. different from that in the Temperate and Torrid Zones. Light is a primary synchronizer for biological rhythms. At the latitude of Wainwright, direct solar radiation is GROWTH absent 4 months of the year and present 4 months in summer. Anthropometric Measurements Studies were made to determine the effects of this un- Anthropometric measurements will be used in charac- usual light-dark cycle on Eskimo body rhythms. Thirteen terizing Wainwright Eskimos by their size and shape. subjects were studied intensively for a 10-day period be- They will also be used in comparing Eskimos with non- fore and after each of two solstices and two equinoxes. Eskimo groups and in calculating biological distances be- Physiological observations were made and urine was col- tween Eskimos in Alaska, Canada, and Greenland.Data lected every 2 hr while the subjects were awake. Oral from the summer of 1968 will be used as the first inter- temperature, pulse, blood pressure, eye-hand coordina- cept of a longitudinal growth study and willbe used tion, and grip strength were measured. Urine samples cross-sectionally in the same study. Physiologists are were analyzed for volume, pH, potassium, sodium, chlo- using skin-fold measurements for calculating lean body ride, calcium, and the steriods 17-0HCS and 17-KS. The mass. Anthropometric data will also be applied tothe data were analyzed and rhythm parameters computed by study of hybridization. the Periodicity Analysis Laboratories at the University of The following general statements about Wainwright Minnesota. Eskimos are based on anthropometric measurements: Of the data collected during the summer of 1968, Adult females are only 2 lb lighter than adult males, only those pertaining to oral temperature, urine potas- but they are 10.5 cm shorter in stature. sium, and urine chloride were sufficient to show definite Males remain lean and muscular until they are old daily rhythms. Data for the entire year are being ana- and average 6 lb less than Caucasian standards. Females lyzed for animal rhythms. tend to put on weight during adolescence and are about 20 lb heavier than Caucasian standards. Hybrids of both sexes are taller and heavier than OPHTHALMOLOGY pure Eskimos. A secular growth trend for stature was seen in those born around 1880 as comparedwith those An ophthaLmologist examined the eyes of 191 persons born in 1940. for refractive error, iris color, eye disease, color sense, and climatological changes. He made a number of anthro- Bone Mineral Measurements pometric measurements, including (1) thickness and breadth of corneal radius and (2) pupillary distance, and Measurements of the mineral content of bone were he made a number of anthroposcopic observations on the made by a direct photon absorption method. The mono- site of iris frill, the existence of embryotoxon corneae energetic radionuclide photon source was 1251. The

67 shafts of the radii and ulnae of 28 children and 24 adults with perceptual processes. The qualities tested and the were scanned for calcium hydroxyapatite. tests used were as follows: The bone mineral values for the Eskimo sample were Spatial visualization; a revision of the Gilford- as higli as, or higher than, values for U.S. Caucasians of Zimmerman spatial visualization IV tests corresponding age and sex. Both Eskimo children and Ability to identify embedded figures; the embedded adults, however, had a surprisingly high mineral content figures of Witkin relative to their small stature. Two older subjects were Digit span; one of the subtests of the Wechsler osteoporotic, and two others showed signs of diminished Adult Intelligence Scale bone mineral conte. t. Another series was concerned with cognitive processes. The tests used were Raven's Progressive Matrices, Dentition Thurstone's Code Aptitude Test, and tests in English and Inupiak (Eskimo) vocabulary. The population may be divided with respect to dental caries into the following groups: All necessary translations were prepared at Point Bar- Older persons. Their teeth are well worn from at- row before going to Wainwright. Assistance was received trition and abrasion, but with no caries. The teeth of from native Eskimo speakers at the Naval Arctic Research these persons developed before sweets, refined flour, and Laboratory. foods from non-Eskimo sources were readily available. Although data have not been completely analyzed, Middle-aged persons. Forty-three are toothless be- certain findings can be reported. No sex difference was cause of cariogenic European foods. They have had little found in the responses to the spatial visualization test. dental care. A marked sex difference is found in the responses of Cau- Eskimos in schooL They have a very high caries casians to this kind of test. Data from the mental tests rate but are receiving dental care and fluoride treatment show a slow growth rate that continues to age 20. In con- regularly from Public Health Service dentists. trast, growth ceases at age 16 in the United States. In Although there were accumulations of debris with these tests, the performance of a 16-year-old Eskimo is heavy deposits of calculus, these did not produce the in- equivalent to that of a 13-year-old Caucasian in the flammatory conditions that would have occurred in most United States_ other groups. A high percentage of the population is mandibularly prognathic. This causes difficulty with dentures in later GENETICS years and results in orthodontic problems peculiar to Eskimo children, who have a Class II malocclusion even Demography at age 2 or 3. The Wainwright population was 308 in July 1968 X rays on 91 persons were acceptable for a study of 169 maies (median age 18.6) and 139 females (median facial and dental patterns. The study was carried out at age 13.6). There were 67 hybrids, but only three were the Center for Human Growth and Development, Uni- one half non-Eskimo. Hybridization, which occurred versity of Michigan. The program consisted of a specially about 1900, was through admixture with New England- devised measurement system, automatic print-out and based whalers of many racial strains. An average of 10.6 card-punching equipment, and a matched-pair program matching each Eskimo with a U.S. Caucasian of the same children have been born alive to the 17 postmenopausal age and sex. The summary conclusions are as follows: women. There are 201 children in 61 sibships. The popu- The Eskimo mandible is protruded and rotated lation has increased more than 3 percent per year over downward in relation to the facial structure. the last 10 years. Nine sets of twins born alive in 364 The palatal and lingual inclination of the upper and conceptions yielded a twinning incidence of 273 per lower incisors is greater in the Eskimo than in other races. thousand birthsmore than twice the incidence of twin- Changes in facial patterns by aging and the loss of ning in Caucasian populations in the United States. The occlusion are great in the Eskimo. male:female ratio for live births was 108.2:100_ More The males have larger faces and mandibles than the male infants died than females, but female out-migration females. caused the present-day sex ratio favoring males. Only 81 percent of the children born alive have survived beyond age 5. This is in contrast to survival of about 97 percent in the United States. Differential fertility is shown PSYCHOLOGY by the fact that one fifth of the women are responsible for almost one half of the living offspring. Intentional A battery of psychological tests was administered birth spacing was not practiced until it was introduced by to a group of about 75 persons. One series was concemed the U.S. Public Health Service a few years before this

68 "70 study was begun. A high infant mortality was the only The psychiatrist working in this program found four population regulator before introduction of medically persons at Wainwright with recurrentpsychopathologies. supervised contraceptive techniques. These wiil be screened for EEG abnormalities at the labo- Pedigree analysis, extending from the present through ratory of the Institute of Arctic Biology in Fairbanks.In the ascending fourth generation, revealed that 4 outof addition, serum-bound calcium levels will be followed in 32 marriages in the parental generation are consanguin- a search for evidence of correlationsbetween hypocal- eous. These four marriages are: onebetween second cemia and mental disorder. cousins, one between second cousins once removed, and two between first cousins once removed.Sewell Wright's mean coefficient of relationship for the parentsis 0.005, a value lying between third cousins andthird cousins APPENDIX A: PUBLICATIONS once removed. Bohlen, J. G. 1970. Circadi- and circannual rhythms in Wain- Wright's coefficient of inbreeding was calculated. Cal- wright Eskimos. Arctic Anthropol. 7(1):95-100. Feldman, C. F., and R. D. Bock. 1970. Cognitive studies among culations using data from pedigree analysis yielded a residents of Wainwright Village, Alaska. Arctic Anthropol. mean value of 0.004, or equal to third cousins,while data 2:101-108. from MN blood groups yielded a value of 0.19, or a value Jamison, P. L. 1970. Growth of Wainwright Eskimos: stature and weight. Arctic Anthropol. 2:86-94. between that for sibs or uncle-niece. The coefficient cal- Jamison, P. L., and S. Zegura. 1970. An anthropometric study culated from blood group data is obviously too high, and of the Eskimos of Wainwright, Alaska. Arctic AnthropoL pedigree analysis underestimates the consanguinity of the 2:125-143. Mayhall, J. T. 1970. The effect of culture change upon the ancestral population. Eskimo dentition. Arctic Anthropol. 2:117-121. Mayhall, J. T., A. A. Dahlberg, and D. G. Owen. 1970. Torus mandibularus in an Alaskan Eskimo population. Amer. J. Dermataglyphs Phys. Anthropol. 33:57-60. Finger and palm prints were obtained on 97 males and Mazess, R. B. 1970. Bone mineral content in Wainwright Eskimos: a preliminary report. Arctic AnthropoL 2:114-116. 112 females at Wainwright, and on 13 males and 21 fe- Milan, F. A. 1968. The international study of Eskimos. Arctic males at Anaktuvuk Pass. For comparative purposes there 21:123-126. Kodiak Island, Milan, F. A. 1970. Preliminary estimates of inbreeding levels in are prints on 96 Eskimos from Karluk, Wainwright Eskimos. Arctic AnthropoL 2:70-72. and about 120 Eskimos and Indians from Alakaket; these Milan, F. A. 1970. The demography of an Alaskan Eskimo vil- were all made earlier. In addition to patterns(radial, ul- lage. Arctic Anthropol. 2:26-44_ nar, and whorl), palmar prints will be used to screenfor Milan, F. A. 1970. The demography of an Eskimo village on the north slope of Alaska. Arctic 23:82-99. chromosomal aberrations. Karyotype analyses have been Milan, F. A., ed. 1967. Report on the working party conference undertaken on some of these persons, especially those of for the IBP/HA study of circumpolar populations held at Anaktuvuk Pass, by the Battelle Memorial Institute under Point Barrow, Alaska, 17-22 November 1967. University of Wisconsin, Madison. sponsorship by the Atomic Energy Commission. Rennie, D. W., P. DiPrampero, R. W. Fitts, and L. Sinclair. 1970. Physical fitness and respiratory function of Eskimos of Wainwright, Alaska. Arctic AnthropoL 2:73-82. Saliva Analysis Robinhold, D., and D. Rice. 1970. Cardiovascular health of Two hundred twenty-one saliva specimens were ana- Wainwright Eskimos. Arctic Anthropol. 2:83-85. Sauberlich, H. E., W. Goad, Y. F. Herman, F. A. Milan, and lyzed for blood group substance. All Eskimos secreted P. L. Jamison. 1970. Preliminary report on the nutrition blood group substances in the saliva; 55.6 percent were survey conducted among the Eskimos ofWainwright, Alaska, A, 31.7 percent were 0, 7.7 percent were B, and5_0 per- 21-27 January 1967. Arctic Anthropol. 2:122-124. Way, A. B. 1970. A method of measuring general health and its cent were AB. relationship to effective fertility in Wainwright Eskimos. Arctic Antliropol. 2:109-113.

BEHAVIOR APPENDIX B: PARTICIPANTS The clinical psychologist of the Alaska Native Health Frederick A. Milan, Arctic Health Research Center, US. Public Service has described two related sociopathologies that Health Service, College, Alaska (Program Director; demog- are constantly seen in working withEskimos. The first raphy; reproduction and inbreeding) consists of numerous manifestations of physical violence, Fred Allen, ir., The New York Blood Center, New York, N.Y. (blood group genetics) particularly those of a self-destructive sort, such as sui- T. Bates, Department of Radiology, Medical School,University cide, accidents, or child abuse. The second consists of of Oregon, Portland (medicine) various failures of communication or verbalization, and R. Darrell Bock, Department of Education, University of Chi- cago, Chicago, IIL (psychology) the substitution of action for words. This leads to a dis- B. Boettcher, School of Biological Sciences, The Flinders Uni- associative process in which the connection between versity of South Australia, Bedford Park, South Australia meaning and behavior is lost. (salivary amylase)

69 J. G. Bohlen, NIH Predoctoral Fellow, Human Biology Program, J. Mayhall, Zoller Memorial Dental Clinic, University of Chi- University of Wisconsin, Madison, and Department of Path- cago, Chicago, Ill. (dental health and growth) ology, University of Minnesota Medical School, Minneapolis Richard Mazess, Department of Radiobiology, University of (chronobiology) Wisconsin, Madison (anthropology) D. Bosman, University of Wisconsin, Madison (preventive med- D. McLean, University of Chicago (psychology) icine) Robert J. Meier, Department of Anthropology, Indiana Univer- J. BrOsted, University of Copenhagen and University of Wiscon- sity, Bloomington (anthropology) sin (anthropology) Mary Jane Moore, Graduate Student, Department of Anthro- A. A. Dahlberg, Director, Zoller Memorial Dental Clinic, Uni- pology, University of Wisconsin, Madison (carbonic anhy- versity of Chicago, Chicago, Ill. (dental health and growth) drase) Mrs. A. A. Dahlberg, Project Assistant, Zoller Memorial Dental Richard H. Osborne, Professor of Anthropology and Medical Clinic, University of Chicago, Chicago, Ill. (dental health and Genetics, University of Wisconsin, Madison (blood group growth) genetics) P. DiPrampero, Instituto Di Fisiologia Umana, Milan, Italy D. Owen, Zoller Wmorial Dental Clinic, University of Chicago, (work physiology) Chicago, Ill. (dental health and growth) Charles T. Dotter, Department of Radiology, Medical School, Frank Pau Is, Section of Laboratories, Division of Public Health, University of Oregon, Portland (medicine) Pouch H., Juneau, Alaska (epidemiology) Harold H. Draper, Professor of Nutritional Biochemistry, Uni- Jack E. Pegg, Department of Radiology, University of Oregon, versity of Illinois, Urbana (nutrition) Portland (medicine) Carol S. Feldman, Department of Psychology, University of Donald W. Rennie, Department of Physiology, State University Chicago, Chicago, Ill. (psychology) of New York, Buffalo (work physiology) R. W. Fitts, Graduate Student, Biodynamics Laboratory, Uni- D. B. Rice, Fredricksburg, Va. (medicine) versity of Wisconsin, Madison (work physiology) D. Robinhold, Resident in Internal Medicine, University of Henrik Forsius, Oulun Yliopisto, Silmitautien Klinikka, Univer- Colorado Medical Center, Denver (medicine) sity Eye Hospital, Uuskikatu 50, Oulu, Finland (medicine) Howard E. Sauberlich, U.S. Army Medical Research and Nutri- Edward F. Foulks, Consultant Psychiatrist, Department of tional Laboratory, Fitzsimons General Hospital, Denver, Health and Welfare, Fairbanks, Alaska (psychiatry) Colo. (nutrition) W. Goad, U.S. Army Medical Research and Nutrition labora- Lynn Sinclair, Graduate Student, Department of Physiology, tory, Fitzsimons General Hospital, Denver, Colo. (nutrition) State University of New York, Buffalo (work physiology) Franz Halberg, Department of Pathology, Chronobiology Labo- W. Thompson, Chief, Arborvirus Research Unit, Department of ratories, University of Minnesota Medical School, Minneap- Preventive Medicine, University of Wisconsin, Madison (epi- olis (chronobiology) demiology) P. Hogan, State University of New York, Buffalo (physiology) Anthony Way, NIH Postdoctoral Fellow, Department of Medi- P. L. Jarnison, NIH Predoctoral Fellow, Department of Anthro- cine, and Graduate Student, Human Biology Program, Uni- pology, University of Wisconsin, Madison (anthropology) versity of Wisconsin, Madison (medicine) Solomon H. Katz, Department of Anthropology, University of D. Wilson, State University of New York, Buffalo (physiology) Pennsylvania, Philadelphia (medicine) M. Wilson, State University of New York, Buffalo (physiology) 0. Lee Kline, American Institute of Nutrition, Bethesda, Md. D. Wishart, Department of Radiology, University of Oregon (nutrition) Medical School, Portland (medicine) W. S. Laughlin, Department of Anthropology and Biobehavioral S. Zegura, NIH Predoctoral Fellow, Department of Anthropol- Sciences, University of Connecticut, Storrs (anthropology) ogy, University of Wisconsin, Madison (anthropology) T. Lewin, University of Gothenburg, Sweden (anatomy)

70 I IM Air lE CO noseAnumam PR() da IFt A MI S

73 ENVIRONMENTAL COMPONENT Biology and Ecologyof Nitrogen

Research into the biology and ecology of nitrogen, trogen, and (3) managing fixed nitrogen in the environ- which is a critical phase of the worldwide effort to pro- ment. Investigations concerning some of these matters vide food and nitrogen for an expanding population, is are under way. conduc red in accordance with unusual requirements. It is probable that the methods for providing food and nitrogen a hundred years from now will differ radically OBJECTIVES from today's methods. For the next 50 years or so, however, it will be necessary to depend primarily onmethods now available. The main business ofresearch will be to The objectives of the program are as follows: refine and expand those methods. To make direct observations on the fixation of ni- Nitrogen is one of the inputs necessary for producing trogen in representative soils, giving special attention to food and, at the same time, is one of the more likely agricultural soils sources of environmental degradation. Thus, thereis a To examine oxygen and nitrous oxide concentra- great demand for increasing nitrogen utilization in ways tions in deep soil profiles to obtain information en de- that will not degrade the environment. nitrification, paying particular attention to agricultural There are indications that intensive agricultural pro- ecosystems where intensive inputs of fixed nitrogen and duction by methods followed in many parts of the world other fertilizers are common today could meet anticipated needs for food and protein To determine nitrate concentrations in deep pro- production. On some of our more productive lands, these files of arid soils, paying particular attention to soils that methods entail the input of nitrogen fertilizer at rates are expected to come under irrigation in the nearfuture 50 to 100 times greater than the nitrogen input rates of and to areas where the presence of underlying concentra- native ecosystems. As production potential is maximized, tions of nitrates might be inferred from geologic evidence even heavier applications of nitrogenous fertilizers canbe To organize conferences of workers interested in anticipated. In many cases the nitrogen harvested with a problems of nitrogen management in agricultural ecosys- crop amounts to no more than 20 to 30 percentof that tems, including the problem of dispoing of nitrogenous applied, and the remaining nitrogen, usually in the form wastes of nitrate, becomes a potential health hazard in foods To publish and circulate abstracts of work in prog- and feeds or in streams and aquifers. As such it can be a ress dealing with problems of nitrogencycling and reac- strong contributor to eutrophication. tions of the nitrogen cycle The logistics and economics of intensive agriculture To coordinate related investigations of nitrogen tend to favor the input of industrially fixed nitrogen over movement in native ecosystems in the analysis-of- the use of nitrogenous organic wastes for fertilizer, and ecosystems program the wastes become an additional potential pollutant of the environment. Means for disposing of these wastes must be developed. A conference on the biology and ecology of nitrogen RESEARCH PLANS was held at Davis, California, inNovember 1967 under the auspices of the U.S./IBP. Discussions were concerned chiefly with (1) the need for a more detailed understand- This program includes research on problems of nitro- ing of the inputs and outputs of fixed nitrogen both in gen management that are not included inother U.S./IBP native ecosystems and in agricultural systems, (2) means programs, and it integrates information on nitrogen that of efficiently providing adequate quantities of fixed ni- is being developed in other programs.

73 RESEARCH TOPICS tilized fields. Nitrous oxide is a good presumptive indicator of denitrification activity, and oxygen levels will Nitrogen Fixation indicate the likelihood of denitrification. Denitrification probably is not significant at oxygen tensions above Nitrogen fixation by free-living organisms and by 10 mm. Where possible, nitrate levels will be determined symbiotic systems will be evaluated. One procedure will in profile to aid in interpretation. be the direct evaluation of fixation by using isotopic nitrogen and by applying the acetylene-ethylene reaction. Work to date has suggested that the acetylene-ethylene Nitrate Determination in Deep Profile reaction is exceedingly useful as a means of evaluating Work has been done in determining salt content in nitrogen-fixing capabilities, but the calibration between deep profile. Because of difficulties experienced in de- this technique and information gained by direct isotopic termining nitrate content, most of this work has not pro- means is variable. The two methods must be used in con- vided information regarding nitrate. Future studies will junction if reliable information is to be obtained. Isotope- require prompt analysis of samples before nitrate losses dilution methods can also be employed, but the cost of occur. isotopic nitrogen limits this technique. Should isotopic nitrogen become less expensive, the dilution technique will be used more often. COORDINATION AND MANAGEMENT

Denitrification Conferences Where possible, rates of denitrification will be mea- Several conferences are planned in which 15 to 20 sured directly by isotope-dilution methods. In most cases, workers will be brought together to exchange informa- however, the significance of denitrification will have to tion and to participate in workshops. They will be held be evaluated by indirect methods and by inference from in areas where nitrogen problems are likely to be acute, oxygen and N20 data. including the Midwest (intensively managed soils), the Southwest (arid areas near population centers), and urban Nitrates in Groundwaters areas where domestic wastes and the intensive use of ni- The delivery of nitrogenous compounds to ground- trogenous fertilizers may contribute to the contamination of water. waters and to bodies of freshwater will be monitored by direct measurement and by collating data already available. Information Exchange A newsletter and abstracts of research results will be Oxygen and Nitrous Oxide Determinations distributed from the University of California, Davis. Dis- tribution will be made initially to laboratories where The oxygen and nitrous oxide status of soils in deep profiles will be examined. Sampling will be accomplished workers are involved in investigations relate-1 to nitrogen. p&narily by means of capillary plastic tubing connected Information for the newsletter will be contributed by the workers. with equilibration reservoirs at various depths in profile. This method has been selected in preference to direct Management Personnel oxygen electrode determinations because the calibration of the electrodes are unstable with time. Gases from im- The Program Director is C. C. Delwiche, Professor of planted reservoirs can be sampled periodically by a com- Geobiology, Department of Soils and Plant Nutrition, bination of gas chromatographic, infrared, and mass University of California, Davis. When funded, the pro- spectrometric measurements. Areas selected for this kind gram will be guided by an executive committee consisting of analysis will include heavily irrigated and heavily fer- of participating research workers.

74 Biological Control ofInsect Pests

The purpose of the program on biological controlof OBJECTIVE insect pests is to study all principles ofbiological control of thc en- that can be applied without undue disturbance The general objective of this program is toutilize ali vironment. The following principles are amongtho: that methods of pest control to develop the scientific basis will be studied: for an integrated method of controlling populationsof Autocidal techniques insect pests at non-economic densities in a mannerthat Biochemical regulation of development and repro- optimizes cost-benefit relations and minimizes environ- duction Pheromones, attractants, repellents, and barriers mental degradation. Population dynamics Plant resistance to attack by insects The potential of autocidal techniques was shown RESEARCH PLANS dramatically by a program in which the Agricultural Re- search Service (ARS) eradicated the screw-wormfly from the southeastern part of the United States. This program has two phases. The first is pest-group The potential of biochemical regulators (hormones)is centered and embraces the use of natural enemies.It is suggested by the existence in certain phylogenetically part of an international effort tofind better biological ancient plants (e.g., ferns and horsetails) of chemicals means of controlling insects.It is supported by ARS, by that interfere with insect development and by the fact state agricultural experiment stations, aridby universities. that few insect species have adapted to feed on these About 30 projects are included in this phase, andthey plants. The chemicals are similar to insect hormones. are concerned with four groupsof pests: (1) aphids, The potential of biochemical messengers (pheromones) (2) rice pests and th e. cereal leaf beetle, (3) spider mites, is suggested by results obtained in trapping studies in and (4) scale insects. which natural and synthetic pheromones were used. Oc- The second phase is concerned with a newer,broader currence of two or more chemicalstogether can present effort: integrated control. In this type of control, all either a blockage or a synergistic (additive) effect.The compatible measures are used, but the use of pesticides complexities of the interaction of the chemicals need ex- is minimized. The work will be site-centered and crop- tensive study. The effect of the chemicals on natural ene- centered. The major effort will be made at the popula- mies of the target species also need study. tion level in agroecosystems. Only in this way can weob- The role of population dynamics is demonstratedby tain the information necessary for modeling andfor the control of insect pests by natural enemies, such as predicting consequences of control measures. predators, parasites, or pathogens. For many pest species The Program Director is C. B. Huffaker. there exist somewhere (often right in the same environment) species of natural enemies that are capable of controlling pest densities at levels below those at whichseri- ous crop losses result. Whereeffective enemy species are RESEARCH METHODS AND PROGRESS deterred from exerting their potential roles, or do not exist, enemy species can be introduced. On aworldwide the basis, there have been about 250 instances of successful Research progress thus far has been limited to introduction of enemy species. About one third ofthese older of the two phases of the biological control pro- gramthe internationally organized phase that is con- are so -uccessful that the threateningspecies present no problem at all. cerned with four groups of pests.

75 APHIDS APPENDIX A: PUBLICATIONS DeBach, P., and C. B. Huffaker. In press. Experimental tech- The outstanding development in the aphids project is niques for evaluation of the effectiveness of natural enemies, the establishment of a parasite, Trioxys pallidus, from ch. 5.In C. B. Huffaker led.) Biological control. Plenum Iran that has proved highly effective in reducing popula- Press, New York. DeBach, P., D. Rosen, and C. E. Kennett. In press. Biologial tions of the walnut aphid in California. if expectations control of coddids by introduced insects, ch. 7.In C. B. are fulfilled, complete biological control can be achieved. Huffaker ed.] Biological control. Plenum Press, New York. Flaherty, D. L. 1969. Ecosystem trophic complexity and Workers in Minnesota have discovered sources of wild Willamette mite, Eotetranychus willamenei Ewing (Acarina: potato germ plasm that is resistant to the aphid Mysus Tetranychidae), densities. Ecology 50:911-91 6. persicae. Hybrids from crosses with cultivated potato are Flaherty, D. L.. and C. B. Huffaker. In press. Biological control of Pacific mites and Willamette mites in San Joaquin Valley being tested in the field by E. B. Radcliffe and F. I. Lauer. vineyards. I. Role of Metaseiulus occidentalis. Hilgardia. Workers in Washington have reported that naturalene- Hagen, K. S., E. F. Sawall, Jr., and R. L. Tassan. In press. The mies of a pest can reduce the population of the pest by use of food sprays to increase effectiveness of entomophagous insects. Proceedings, Tall Timbers Conference on Ecological harassment as well as by parasitization. (See Tarnaki et al., Animal Control and Habitat Management, Tallahassee, Fla. 1970, in the appended list of publications.) Hagen, K. S., and R. van den Bosch. 1968. Impact of pathogens, parasites and predators on aphids. Annu. Rev. Entomol. 13: 325-384. Hoyt, S. G., and L. E. Caltagirone. In press. The developingpro- RICE STEM BORERS AND grams of integrated control of pests of apples in Washington THE CEREAL LEAF BEETLE and peaches in California, ch. 18.In C. B. Huffaker [ed.] Biological control. Plenum Press, New York. Huffaker, C. B., M. van de Vrie, and J. A. McMurtry. 1969. The In the United States, research concerning this group ecology of tetranychid mites and their natural control. Annu. of insects has been concentrated on the cereal leaf beetle. Rev. Entornol. 14:125-174. An effort is being made to establish parasites and to de- Laing, J. E., and C. B. Huffaker. 1969. Comparative studies of predation by Phytoseiulus persimilis Athias-Henriot and velop a computer-oriented program for evaluating their Metaseiulus occidentalis (Nesbitt) (Acarina: Phytoseiidae)on effectiveness. populations of Tetranychus urticae Koch (Acarina: Tetrany- chidae). Res. Pop. Ecol. 11:105-126. McMurtry, J. A., C. B. Huffaker, and M. van de Vrie. Inpress. The ecology of tetranychid mites and their natural control. SPIDER MITES I. The kinds of spider mite enemies, the effects ofspray practices on them and their biological properties. Hilgardia. McMurtry, J. A., and G. T. Scriven. 1968. Studies on predator- Development of integrated control programs offer prey interactions between Amblyseius hibisci and Oligony- possibilities of considerably reducing the use of pesticides chus punicae: effects of host-plant conditioning and limited quantities of alternate food. Ann. Entomol. Soc. Amer. 61: against spider mites on grar.z.s, strawberries, and peaches 393-97. in California and against apples in Washington. In Wash- Tamaki, G., J. E. Halfhill, and D. 0. Hathaway. 1970. Dispersal ington the use of pesticides has been reduced about 50 and reduction of colonies of pea aphids by A phidius smithi (Hymenoptera: Aphidiidae). J. Econ. Entomol. 63:973-80. percent as a result of the modified program. Tamaki, G., J. E. Halfhill, and J. C. Maitlen. 1969. The influence Research in California has shown that spider mite in- of WC-21149 and the aphidiid parasite Aphidius smithiion festations on grapes are due largely to theuse of pesti- populations of the pea aphid. J. Econ. Entomol. 62:678-82. van den Bosch, R., B. D. Frazer, C. S. Davis, P. S. Messenger, cides. Cessation of treatments in experimental vineyards and R. C. Horn. In press. An effective -talnut aphid parasite has been followed by restoration of satisfactory biologi- from Iran. Calif. Agr. cal control. Sometimes, however, full restoration is delayed 3 or 4 years as a result of lingering effects of the APPENDIX B: PROJECTS AND PRINCIPAL pesticides on the predators. INVESTIGATORS* A list of key predators that are effective against key spider mites in the United States was developed. Similar Project: Aphids lists for other countries are also available. G. W. Angalet, Agricultural Research Service, U.S. Department of Agriculture, P.O. Box 150, Moorestown, N.J. (the role of parasites, predators, and disease in the biological control of the pea aphid in the eastern United States) SCALE INSECTS H. C. Chiang, Department Gf Entomology, Fisheries, and Wild- life, University of Minnesota, St. Paul (ecology of predatory insects; trophic relationships of insects found in sugar beet Taxonomic studies to untangle some of the complexi- fields; trophic relationships of insects found in the son in ties of parasites of scale insects have hadsome success, cornfields) and various parasites have been exchanged between the United States and other countries. Biological control of *This list is limited to principal investigators in the phaseof the the California red scale has been improved. program that is concerned with four groups of pests.

76 biological con- K. S. Hagen, Division of Biological Control, University ofCali- spray programs and cultural practices on the trol of mites on apples in Washington) fornia, Berkeley (evaluation of the effectiveness of native of natural enemies of the spotted alfalfa aphid ana other C. B. Huffaker, Division of Biological Control, University aphids; evaluation and augmentation of biological control California, Berkeley (basic studies on the dynamics of agents to -4:place or supplement the use of pesticides to con- arthropod populations; the role of natural enemies in suppressing spider mites in important food crop ecosystems) trol phytophagous arthropod pests) of Cali- J. E. Halfhill, Agricultural Research Service, U.S.Department of C. E. Kennett, Division of Biological Control, University Agriculture, 3706 West Nob Hill, Yakima, Wash. (biological fornia, Berkeley (a study of biological control potentials and control of the pea aphid, A cyrthosiphon pisum (Harris), performance features of predators of spider mites in citrus through mass propagation and release of its braconid para- and deciduous fruit orchards; the biology, ecology, and sys- sites) tematics of the phytoseiidae in California) Florian I. Lauer, Department of Horticultural Science, Univer- J. E. Laing, Division of Biological Control, University of Cali- sity of Minnesota, St. Paul (population dynamics of the green fornia, Berkeley (basic studies on the dynamics of arthropod peach aphid, Myzus persicae, on potato) populations; the role of natural enemies in suppressing P. S. Messenger, Division of Biological Control, University of spider mites in important food crop ecosystems) California, Berkeley (bioclimatic studies with aphid para- C. W. McCoy, Agricultural Research Service, US.Department of sites) Agriculture, Orlando, Fla. (biological control of citrus in- E. B. Radcliffe, Department of Entomology, Fisheries and Wild- sects and mites) life, University of Minnesota, SL Paul (population dynamics J. A. McMurtry, Division of Biological Control, Universityof and of the green peach aphid, Myzus persicae, on potato) California, Riverside (biological control of mites in citrus R. B. Thurston, Department of Entomology,University of Ken- avocado) tucky, Lexington (population dynamcis of the greenpeach M. H. Muma, Department of Entomology, Universityof Florida aphid, Myzus persicae. on potato) Citrus Experiment Station, Orlando (spider mite pathogen R. van den Bosch, Division of Biological Control, Universityof study; natural control of six-spotted mites; biology of California, Berkeley (biological control of pest aphids byim- phytosefidae) E. R. Oatrnan, Division of Biological Control,University of Cali- ported aphidophagous insects) fornia, Riverside (biological control of the two-spotted spider mite on strawberry) A. G. Selhime, Agricultural Research Service, U.S.Department Project: Rice Pests of Agriculture, Orlando, Fla. (biological control ofcitrus in- H. A. Bess, Department of Entomology, Universityof Hawaii, sects ano mites,` Honolulu (biological control of rice pests) D. L. Haynes, Department of Entomology, MichiganState Uni- versity, East Lansing (influence of biotic regulatory agents Project: Scale Insects leaf beetle, Oulema on the population dynamics of the cereal L. E. Caltagirone, Division of Biological Control,University of melanoplus) California, Berkeley (biological control of the olivescale, the K. Nishida, Department of Entomology, University ofHawaii, San Jose scale, and other armored scales in northernCali- Honolulu (biological control of rice pests) fornia) J. D. Paschke, Department of Entomology, Purdue University, H. A. Dean, Department of Entomology, WeslacoAgricultural Lafayette, lnd. (biological control of the cereal leaf beetle, Experiment Station, Texas A & M University, Weslaco (con- Oulema melanoplus) integra- Univer- trol of various mite and insect pests of citrus through F. IV. Stehr, Department of Entomology, Michigan State tion of chemical and biological programs) sity, East Lansing (influence of biotic regulatory agents on P. DeBach, Division of Biological Control, University ofCali- the population dynamics of the cereal leaf beetle, Oulema fornia, Riverside (the role of entomophagous insects in popu- melanoplus) lation regulation of armored scale insects (Hymenoptera: Diaspididae) in major world ecosystems; biosystematics and phylogeny of species of Aphytis of the world; biological con- Project: Spider Mites trol of red, yellow, purple, and other diaspine scales on Entomol- citrus, avocado, walnut, and ornamentals) D. Asquith, Fruit Research Laboratory, Department of C. B. Huffaker, Division of Biological Control, Universityof ogy, Pennsylvania Stat.: University, Arendtsville(basic re- pesticide- California, Berkeley (biological control of the olive scale, the search on the suppression of orchard mites with San Jose scale, and other armored scales in northernCali- resistant predators) of fornia) L. E. Caltagirone, Division of Biological Control, University of Cali- California, Berkeley (a study of biological control potentials C. E. Kennett, Division of Biological Control, University in fornia, Berkeley (biological contol of the olive scale,the and performance features of predators of spider mites Cali- citrus and deciduous fruit orchards) San Jose scale, and other armored scales in northern H. A. Dean, Department of Entomology, Weslaco Apicultural fornia) C. W. McCoy, Agricultural Research Service, US. Department Experiment Station, Texas A & M University, Weslaco (con- citrus in- trol of various mite and insect pests of citrus throughintegra- of Agriculture, Orlando, Fla. (biological control of sects and mites) tion of chemical and biological programs) Mississippi D. L. Flaherty, Division of Biological Control, Universityof M. F. Schuster, cjo Department of Entomology, State University, College Station (biological control of the California, Berkeley (the role of natural enemies in suppress- of ing spider mites in important food crop ecosystems) Rhodes grass scale, Antonina grarninis, through use S. G. Hoyt, Department of Entomology, WashingtonState Uni- Neodusmetia sangwani hi Texas) A. G. Selhime, Agricultural Research Service,U.S. Department versity, Wenatchee (the effect of short feeding periods and in- feeding position of Typhlodromus occidentalis on mortality of Agriculture, Orlando, Fla. (biological control of citrus and fecundity of Tetranychus mcdanielt the influence of sects and mites)

77 Aerobiology

Aerobiology is the study of biologically significant OBJECTIVES materials in the atmosphere. These include spores, pollen, pathogenic microbes, minute animals (suchas aphids), and noxious gases and other pollutants affecting plants, The objectives of the program are: animals, and man. The aerobiology program promotes To promote and coordinate internationally ori- multidisciplinary studies that provide an understanding ented research projects in criticalareas of aerobiology. of how the atmospheric transport of biological matenals To develop improved observation techniques and affect ecological systems. In view of the goals of the IBP, data standardization that will permit comparison ofrec- the scope of the aerobiology program has been broadened ords and information handling systemson an interna- to include the evolutionary development of airborne bio- tional basis. logical particles as well as productivity of plant ecosys- To increase and improve exchange of information tems under the stress of disease caused by airborne patho- between discipiines and between nations. gens. The IBP provides a unique opportunity to investigate these aerobioiogy phenomena ona global basis. Aerobiology was accorded a place in the early plans of section UM. The U.S./IBP effort began with planning in RESEARCH DESIGN 1966 and 1967 and was soon augmented by the IBPpro- gram of the American Phytopathological Society. In April 1969 the aerobiology program received funds for management and general activities and about 1 month The program consists of eight discipline-orientedseg- later opened a program office in Ann Arbor, Michigan, ments representing areas where information is urgently and a liaison office in Beltsville, Maryland. Theprogram needed: spore loads in the atmosphere, evolutionof air- plan was discussed by the American Phytopathological borne pathogens, plant production losses dueto airborne Society at the August 1969 meeting in Spokane, Wash- pathogens, allergenic particles in the atmosphere, phyto- ington, and by members of the 12th International Bo- geography and genecology of pollen andspores in the tanical Congress in Seattle later in thesame month. atmosphere, human and animal pathogens in theatmo- An advisory committee, established to guide policy sphere, insects and other microfauna in the atmosphere, and to review research proposals, held its firstmeeting in and effects of gaseous and particulate pollutantson Beltsville on November 20 and 21, 1969. Officers of the plants and animals. The first three segmentsconstitute phytopathology subprogram met in Washington, D.C., the phytopathology subprogram. January 28 and 29, 1970, to revise segment objectives William S. Benninghoff, Department of Botany,Uni- and to consider possibilities for funding researchpropos- versity of Michigan, Ann Arbor, is Program Director. als. A workshop on plant production losses dueto disease W. D. McClellan, Crops Research Division,Agricultural was held in St. Paul, Minnesota, February 26-28, 1970. Research Service, U.S. Department of Agriculture, Belts- It was attended by representatives of variousagencies and ville, Maryland, is director of the phytopathologysub- institutions of Canada and the United States. Theadvis- program. Segment supervisors are as follows: ory committee held its second meetingon June 2 and 5, 1970, in Raleigh, North Carolina. This coincided withthe Spore loads in the atmosphere: Richard D. Schein, Col- Raleigh conference described under "ResearchProgress," lege of Science, Pennsylvania State University,Uni- below. On June 3 the committee met with theIBP Aero- versity Park biology Theme International Working Group,which had Evolution of airborne pathogens: William Snyder, Depart- been convened on the same occasion. ment of Pathology, University of California, Berkeley Plant production losses due to airborne pathogens: Lucas Calpouzos, Department of Plant Pathology, University of Minnesota, St. Paul Allergenic particles in the atmosphere: William Solomon, tives in Atmospheric Monitoring," was held at Raleigh, University of Michigan Medical Center, Ann Arbor North Carolina, in June 1970 and involved 65 persons Phytogeography and genecology of pollen and spores in from 5 countries, 10 government agencies, and 11 univer- the atmosphere: William S. Benninghoff (seeabove) sities. This conference was sponsored by the aerobiology Human and animal pathogens in the atmosphere: H.F. program and the National Air PollutionControl Adminis- Hasenclever, National Institute of Allergy and Infec- tration. The chief accomplishments were: (1) a reviewof tious Diseases, National Institutes of Health,Bethesda, the current status of atmospheric monitoring programs, Md. (2) identification of needed aerobiological information, Insects and other microfauna in the atmosphere: J. and (3) development of recommendations for improving Linsley Gressitt, Bernice P. Bishop Museum, Honolulu, present monitoring systems. A report on theconference Hawaii is being prepared for publication. Effects of gaseous and particulate pollutants on plants and animals: Howard E. Heggestad, Plant Air Pollution Laboratory, Agricultural Research Service, U.S. De- SPORE LOADS IN THE ATMOSPHERE partment of Agriculture, Beltsville, Md. This segnent is concerned with dissemination of plant A program coordinator will be appointed to assistthe pathogens. The research is concentrated on release, trans- Program Director. port, and viability of propagules. An advisory committee is responsible for guiding the program and for evaluating researchproposals. F. A. Air Spora in Hawaii Wood, Department of Plant Patholou, Pennsylvania Diurnal and seasonal periodicities of airborne fiingi State University, University Park, is chairman. ThePro- have been recorded in papaya, macadamia, and banana gram Director, the director of thephytopathology sub- plantations in Hawaii. Periodicities have been related to program, and the segment supervisors aremembers. specific plant diseases, nature of vegetation in areas sam- Other members are as follows: pled, and meteorological conditions. Preliminary studies have been made on abundance and fluctuations in num- Lester Machta, National Atmospheric and Oceanographic bers of possible allergens in residential areas. Cooperation Administration, U.S. Department of Commerce, Silver has been established with the Department of Agriculture, Spring, Md. Fiji, for comparison of airborne particles in banana plan- Donald S. Meredith, Department of Plant Pathology, tations in Fiji and Hawaii. University of Hawaii, Honolulu Donald H. Pack (alternate for Dr. Machta), National At- Supported by National Institutes of Health. mospheric and Oceanographic Administration, U.S. Department of Commerce, Silver Spring, Md. Principal investigators: Ivan W. Buddenhagen and Donald Carl M. Shy (alternate for Dr. Thompson), National Air S. Meredith, Department of Plant Pathology, Univer- Pollution Control Administration, U.S. Department of sity of Hawaii, Honolulu Health, _;cation, and Welfare, Durham, N.C. Jack E. Thompson, National Air Pollution Control Ad- Inoculum Production, Dissemination, and ministration, U.S. Department of Health, Education, Infection by Scirrhia (Dothistroma) pini and Welfare, Durham, N.C. Submitted for funding.

Principal investigator: Fields W. Cobb, Jr., Department of Plant Pathology, University of California, Berkeley RESEARCH PROGRESS The Relationship of Origin and Variability to Pathogenicity and Morphology of Fusarium The research progress of the aerobiology program con- roseum sisted of one program-wide conference and of activities associated with the program segments described below. Work on this project is under way. Arrangements have As a service to the researchers in the program, an aerobi- been made for cooperative work on Fusaria in Australia. ology newsletter was established and is being mailed to about 350 scientists in the United States and nine foreign Submitted for funding. countries_ The program-wide conference, "Aerobiology Objec- Principal investigator: Paul E. Nelson, Department of

80 Plant Pathology, Pennsylvania State University, Uni- been developed, and the investigators have worked with versitY Pa rk the Waite Agricultural Research Institute in South Aus- tralia in perfecting the techniques. Commitments for cooperation have been obtained from Iran, Chile, Central EVOLUTION OF AIRBORNE PATHOGENS America, Malaysia, Japan, Puerto Rico, and a number of U.S. and European workers. This segilIentis concerned with investigation on a world basis of theoriginof plant pathogens and diseases, Submitted for funding. the origm and distribution of inoculum and genes for and the evolution of pathogenicity. Principal investigators: John R. Parmeter, Jr., and A. R. Weinhold, Department of Phytopathology, University Origin and spread of Airborne of California, Berkeley Inoculum cfFusariumPathogens Origin and Distribution of Species Related studieson Fusarium ecology indicate that the of Phytophthora perithecial (irbome) stages of Fusarium species occur Phyrophthora cultures have been obtained from West in certainregions. To date, we have established Africa, Mexico, Guatemala, Costa Rica, and Panama, and major sources for three species, as follows: probable some of these have been tested for compatibility type Gthberella (Fusarium)moniliformeSouth Pacific and for variation in pathogenicity. Contacts for collabo- Caloneetria (Fusariurn)rigiduiscalaAmerican tropks ration have been established in Ghana, Nigeria, Ivoly Coast, France, Brazil, Trinidad, Puerto Rico, Colombia, gYPo'hyces (Fusariurn) solaniJapan and Africa Panama, and Costa Rica. Excellent cooperation has been established with scientists in Asia, Central America, and South America. Submitted for funding. Submitted for funding- Principal investigators: George A. Zentmyer, Donald C. Erwin, and Peter H. Tsao, Department of Plant Pathol- Principal investigators: William C. Snyder, J. G. Hancock, and S. N. $ruith, Department of Plant Pathology, Uni- ogy, University of California, Riverside versitY of California, Berkeley

Species ancl giotypes of the Nematode Genus PLANT PRODUCTION LOSSES DUE MeloidoWne (Root-KnotNematodes) and TO AIRBORNE PATHOGENS CloselY Related Genera The objectives of this segment are to improve methods ThePrincipal investigator visited scientists and admin- of assessing plant production losses due to diseases caused istrators of the International Institute of Tropical Agri- by airborne pathogens. culture In Ihadan, Nigeria, from December 1 to 18, 1969. A workshop on methodology for measuring plant pro- A cooperative research program on the root-knot nema- duction losses due to disease was held February 26 and todes for that part of the world was initiated. In Nigeria 27, 1970, in St. Paul, Minnesota. Twelve scientists from and Peru.the investigator discussed the possibility of a the United States and Canada evaluated the research de- cooperative effort on the part of local institutions and sign, techniques, and interpretations of current work_ North CarOlina State University to enhance professional and subProfftsional competence in nernatology. Yield Losses in Spring Wheat Caused by Puccinia recondite and Puccinia graminis Submitted for tritici in Different Environments A field experiment with nine spring wheat varieties principal investigator: J.N. Sasser, Department of Plant pathology, NorthCarolina State University, Raleigh and four patterns of disease epidemics (stem rust, leaf rust, or both rusts) was carried out in Minnesota, Colo- Origin and bistribution of Rhizoctonia in rado, and Puerto Rico to determine whether there is a Relation to the Evolution and Spread of predictable relationship between disease and loss of yield Pathogenically Specialized Forms in different environments. Data are being analyzed for correlations between severity of disease at different Technique%for identifying pathogenic groupings have stages of wheat development and percentage of loss of

-81 yield. A small workshop on plant losses due todisease eration. Specialists in mycological systematics are being was held in St. Paul, Minnesota,in February 1970. consulted. Communication has been established with scientists in Canada, the United Kingdom, the Netherlands,and Italy Viable Species of Algae and Protozoa regarding the results of this experiment. Negotiations are in the Atmosphere in progress to obtain experimental sites in theNether- Submitted for funding. lands and Yugoslavia. Principal investigator: Harold E. Schlichting, Jr.,-Depart- Supported by Agricultural Research Service, U.S. Depart- ment of Botany, North Carolina State University, ment of Agriculture. Raleigh Principal investigator: Lucas Calpouzos, Department of Plant Pathology, University of Minnesota, St. Paul HUMAN AND ANIMAL PATHOGENS IN THE ATMOSPHERE

ALLERGENIC PARTICLES IN THE This segment is providing liaison between research in ATMOSPHERE the microbiolov-medical mycology area and the aerobiology program_ This segment seeks to evaluate the status of research on airborne allergens, bring ongoingprojects into effective association and stimulate needed research. INSECTS AND OTHER MICROFAUNA There are a number of conditions that make it desir- IN THE ATMOSPHERE able to perform additional research. They include: Lack of pollen concentration data for all but a few This segment is concerned with the atmospheric dis- points in this country and Great Britain persal of minute animals, such as aphids, leafhoppers, The obscurity of the contributions of local pollen and locusts, some of which are vectors of disease. sources (especially trees) to the pollen exposurein any Oven locality Natural Dispersal of Insects in the Arctic Uncertainty regarding the pollen emissions of closely related plant species In the summer of 1969, several thousand specimens Lack of information concerning the effects of of insects and other arthropods were trapped in large ny- ultraviolet light and gaseous atmospheric pollutants on lon nets set out on the tundra near Point Barrow, Alaska, particle allergenicity and some were trapped at Cape Thompson. Additional Lack of information concerrOng the prevalence of trap-nets mounted on a NASA tower near Point Barrow, bacteria, algae, and insect debris in indoor and outdoor set for long-distance dispersants, producedonly a few air specimens. Wind and other weather data are correlated Lack of information on the exposure potential with the trapping results. A paper on these results, and during normal human activity on results obtained by Dr. C. Yoshimotoand assistant in 1965, is in preparation.

Supported by Arctic Institute of North America. PHYTOGEOGRAPHY AND GEN ECOLOGY OF POLLEN AND SPORES IN THE Principal investigators: J. Linsley Gressitt and Eugene ATMOSPHERE Holzapfel, Bernice P. Bishop Museum, Honolulu, Hawaii Several pilot projects are in progress. Donald A. Jameson, Director, Pawnee Site, Grasslands Biome, Studies of Airborne Insects in the Atmosphere analysis-of-ecosystems program, has requested assistance in measuring the import and export of humus by wind Large numbers of insects have been trapped from action at the grassland study sites. Sampling devices and ships in many parts of the Pacific Ocean and in other systems for this problem are being developed. oceans. Cooperation has been receivedfrom individuals Devices that measure concentrations of pollen, espe- and organizations in various other countries, including cially instantaneous concentrations, are being studied. Denmark, England, Chile, and New Zealand. Results of A photographic atlas of fungus spores is under consid- some of this work are being published inPacific Insects.

..,.., :. 2 Supported by Oceanography Branch, Office of Naval Re- This segnent has furnished advice on design of research, U.S. Navy. search projects to predoctoral and postdoctoral investigators and has complied with many requests for informa- Principal investigators: J. Linsley Gressitt and Eugene tion from persons outside this research field. Holzapfel, Bernice P. Bishop Museum, Honolulu, Hawaii Air Pollution Effects on Powdery Mildew Fungi Methods have been developed for producing viable conidia of Microsphaera alni and Erysiphe graminis, for EFFECTS OF GASEOUS AND PARTICULATE transferring conidia directly from hosts to test mem- POLLUTANTS ON PLANTS AND ANIMALS branes with a vacuum spore collector, for even deposition of conidia onto substrates in a settling tower, and for ger- mination under controlled temperature and humidity Objectives: conditions. An ozone exposure apparatus has been de- To collect, on a worldwide basis, information on vised for simultaneously exposing conidia to various levels of total oxidants, sulphur oxides, nitrogen oxides, concentration-time doses of gases and to filtered air. In- fluorides, hydrocarbons, ethylene, lead, cadmium, and vestigations are in progress on the mechanism of induced boron and to determine how the materials were measured resistance of leaf tissue adjacent to mildew colonies when To collect information on losses to vegetation and the tissue is exposed to high concentrations of ozone. animals attributable to air pollutants The objectives are shared with federal and stategov- Supported by National Air Pollution Control Adminis- ernment laboratories, industrial laboratories, and aca- tration. demic laboratories in the United States and other nations. The collected information will be exchanged with other Principal investigator: Craig R. Hibben, Kitchawan Re- nations, and an effort will be made toward uniformity of search Laboratory, Brooklyn Botanic Garden, Os- report procedures and notations. sining, N.Y.

82 Phenology

Man has always been alert to the natural eventsthat Where sufficiently intensive studies have been made, phenological observations may be used for microclimatic take place in his immediate surroundings.In past ages pheno- knowledge of these events was essential to hissurvival. mapping. In the Wingra Lake Basin, Wisconsin, Arboretum Today, even more than in the past, man mustknow, logical data from the University of Wisconsin micro- understand, and record the periodic seasonal changes will be used to make phenological maps indicating climatic conditions affecting productivity in that system. that occur in plants and animals. Thesechanges are col- Microclimatic maps derived from phenological data can lectively termed the phenology of the organisms. moisture The vegetative season varies from year to year,and also be used to indicate the rates at which soil this variation has a profound effect on man andhis abil- is depleted by evaporation. Wherever the responses of plants or animals to heat, ity to produce food. Yet, because so littleinformation light, precipitation, or other factors are observed,pheno- has been accumulated, the variations cannotbe mapped logical data are part of the record for practical and scien- in detail. Data on cultivated crops are availablebut are tific purposes. Examples of applications are given in biased by human manipulation. Table 1.

TABLE 1Examples of Applications for Phenological Data Research Applications Related Sciences Practical Applications

Physical sciences Microcliznatic mapping Climatology Plants as indicators of microclimate Eutrophication trends Limnology Duration of ice-cover, prediction of algal blooms Trafficability, evaporativity, weather Soil science Plants as indicators of soil moisture modification

Agricultural sciences Remote sensing of crops for growth Agronomy Timing of field operations characteristics Prediction of flowering events Apiculture Timing of transplantation of hives Development of adapted varieties Horticulture Timing of propagation Temporal and regional changes in Plant genetics Timing of seed collection and growing season adaptation hereditary traits Pest control through natural enemies Plant patholou Timing of spraying Remote sensing of growth stages and Forestry Timing of Christmas tree shearing, choice of planting sites development of adapted species Life sciences Ecological relationships Investigation of adaptation to Biology environments and evolution Insect development and migration Entomology Pest control Correlation of events with indicator Medicine Prediction of onset of hay fever and allergies produced by plants plants Correlation of development with Pharmacology Timing of collecting plants used in drugs and spices indicator plants

83 OBJECTIVES quiring and retrieving data is an important secondary objective.

The objectives of the phenology program are to develop phenology as a viable science in North America; to INTENSIVE STUDIES elucidate phenological relationships with applications in the physical, agricultural, and life sciences; to assist in Intensive studies of the phenology of particular species integrating IBP research; and to stimulate and coordinate long antedate the IBP, as do studies of the physiological intensive phenological research on mechanisms of pheno- basis of plant and animal responses to phenologically logical response. active stimuli. The phenology program is encouraging additional studies both in intensive investigations of particular species and in the mechanisms of phenological response. RESEARCH PLANS

The phenology program is directed by Forest Steams. INTERNATIONAL COOPERATION The program is in four parts, each of which has a coordinator: Phenological work in the United States is drawing Regional phenological networks (Byron 0. Blair) upon the experience and techniques of networks estab- Standardization of observations and data handling lished in Europe and elsewhere. Efforts are being made (Mrs. Katherine Lettau) to exchange study material between continents and to Intensive studies on specific phenological phenom- assist in establishing cooperative gardens. ena and on mechanisms underlying phenological responses (Richard J. Hopp) International cooperation (Richard J. Hopp) RESEARCH PROGRESS

NETWORKS OBSERVATION NETWORKS Three regional phenological networksnortheastern, north central, and westernare operating in the United To be useful as a phenological indicator, a plant must States. They are funded by cooperative regional projects have certain characteristics: relatively conspicuous and of the state agricultural experiment stations and are co- well-defined developmental phases, timing of develop- ordinated through regional committees. In cooperation ment in tune with agricultural species of value, long life with the northeastern project, phenological observations and adaptation to a wide range of climates, marked re- are now being extended southward. sponse to the meteorological environment, and relative ease of vegetative propagation. The lilac meets these spec- ifications. An additional advantage is that lilacs are estab- STANDARDS lished in a great many locations. Thus, the lilac is a suitable object for phenological observations. Phenological observations must be defined and stan- Honeysuckles of two cultivars were selected to extend dardized if data are to be compared. Standards have been the phenological network in the western region. These developed for lilac and honeysuckle, but much remains cultivars have broad geographic adaptation and are long- to be done for a wide range of other species. Handbooks lived, easily propagated, and relatively free from diseases for observers are planned both for the phenological gar- and insects. They can be observed only 1 year after plant- den network and for a variety of native species. ing. Honeysuckle shows nine distinct phenophases, which A major deficiency in earlier studies was the inability in sequence extend over most of the growing season. to make data readily available to persons wishing to ana- Caprio et al. (1970) describe them as "a precise and ver- lyze them. Some early data were published_ However, a satile climatological instrument." considerable amount of useful data gathered at agricul- At present, studies de7end on observations of lilac in tural experiment stations, U.S. Forest Service labora- the northeastern and north central regions and on obser- tories, and other research organizations is available only vations of honeysuckle and lilac in the western region. in research files. The development of a system for ac- These regional projects are mapping phenologically de-

84 termined environmental zones and examining some of Species growing directly from the soil surface each the mechanisms of phenological response. year are influenced primarily by subsurface soil temperature. Northeastern Region Species with woody structures are influenced by air temperature at screen height and, if they are growing The Vennont Agricultural Experiment Station has as- in heavy soils, by diurnal soil-temperature patterns at sumed leadership in the northeastern network, which in- about 20 inches deep. cludes about 200 sites in 13 states. The dates of several Fifty-five parameters of microclimate of potential phenological events ate recorded each year, and a manual phenological value have been examined statistically, and for phenological observations has been prepared and dis- considerable insight in using analytical techniques has tributed to cooperators (Hopp et al., 1969). been gained. Temperature summations to the dates of phenological In 1968, rye (Secale cereale) was planted at seven sta- events have been calculated for a number of sites. A base tions in Indiana. The performance of this species suggests temperature of 32° F appears to result in a smaller coef- that it may provide data relating directly to the actual ficient of variation than do higher base temperatures. and predicted performance of agricultural crops in vari- January 1 was tentatively used as the starting date for ous parts of the state. temperature accumulation. The results to date are encouraging since they appear to be supported by studies Western Region conducted under controlled conditions. A detailed study related to this project was conducted in New Jersey and In the western United States a strong phenological Michigan in 1969. It resulted in a method for predicting program is functioning under the leadership of Montana harvest dates for blueberries (Addison, 1969). State University. Nineteen publications have been issued, including an exceptionally thorough phenological analy- North Central Region* sis of lilac (Caprio, 1966). The western lilac phenology project includes data from almost 1,000 observation sites In the north central region (12 states), plantings of in 12 states. Results indicate the usefulness of long-term Persian lilac were established at 51 stations. The plantings phenological data for studying the variations in growth were observed at three stages: opening of the first leaf patterns from one year to the next and for examining the bud, full bloom, and end of bloom. These data are being extent of the long-term variability in seasons (Caprio and summarized and evaluated. Metcalf, 1970). Analysis of the data is expected to show In Indiana, the lilac network has been expanded into how variability of weather and of phenological events ob- phenological gardens consisting of 15 to 20 species of served in lilac and certain other species is related to vari- both wild and cultivated plants. Twelve gardens, four of ability of growth in wheat and barley. which have major meteorological facilities, have been es- In addition to lilac, two genotypes of honeysuckle tablished in that state since 1964. Data are available on have been planted at each of 2,000 observation sites. 10 species in 10 locations for 1968 and 1969. One garden Honeysuckle stock is being introduced into the midwest- is functional in Wisconsin. ern and eastern networks. Guides for observing honey- The 12 gardens in Indiana grow shrubs, semishrubs, suckle and lilac have been issued (Caprio, 1968; Hopp et grasses, and perennial forbs. All plants were started by al., 1969; Caprio et aL, 1970). cuttings or by clonal propagation from original sources; Phenological data are available with which to draw de- hence, genetic variation is eliminated. The several species tailed maps showing dates when the purple common lilac provide a flowering range from early April to mid- begins normally to bloom in each of 11 western states. September. Flowering varies from north to south in In- A map for Montana has been published (Caprio, 1966), diana by as much as 3 weeks in the spring-flowering spe- but funds are not available for publishing maps for other cies and a week to 10 days in the fall-flowering species. states. Variations in soil and microclimate are considered. Ten- Cropping hazards are found to be higher in areas in tative conclusions drawn from data of the last several which lilac blooming dates are variable than they are in years are: areas in which the dates are constant.

*On July 1, 1970, the phenology efforts conducted previously Southern Region under the separate regional research projects in the north central and northeastern states were combined and expanded into a new A phenological network has been developed to pro- project (NE 69). This project will include the original programs vide data on several species in North Carolina. Results of described herein and the phenological programs that are being developed in Alaska, Mississippi, North Carolina, and the Cana- early studies have been presented in computer-drawn dian province of Quebec. maps (Lieth and Radford, 1971). Lilac phenology is not

85 86 reliable south of West Virginia, and other species suitable siderably from one another. A relationship between for phenological observations farther south will be se- flushing time and growth rate is indicated by the fact lected. Plants with sharp distinctive phenophases are that late-flushing genotypes show a somewhat faster being sought (Lieth, 1970). Species suitable for aerial ob- growth rate. Knowledge of the variation in flushing dates servation, such as flowering dogwood (Comus florida), and of the genetic makeup of the seedlings is necessary may be extremely useful for correlating remote sensing to determine the degree of genetic control and the rela- data with regional phenological development. tionships between flushing time and temperature summation (Nienstaedt and King, 1969). In central Alberta, seasonal height growth of white INTENSIVE STUDIES spruce seedlings appears to be correlated both with the length of the growing period and with two morphologi- University and government scientists are conducting cal characteristics of the plant: size of the terminal bud phenological studies that differ from the regional net- and height of the seedling (Helium, 1967). The taller the work studies in that they are concerned with plant be- seedling the later the setting of the bud. havior in a small area or with understanding phenological responses. Those described below are representative of Tropical Forest Phenology those included in the U.S./IBP phenology program. In the period 1968-1970, Gordon W. Frankie made observations in Costa Rica on the phenology of several Ecological Effects of Weather Modification hundred mounted specimens of about 160 species of Phenological data and productivity data were collected woody plants that grew in the lowland wet forest at at weekly intervals during the growing season at two sites Finca La Selva, Heredia province, and in the lowland dry 4,700 and 7,800 ft above sea level. Differences in devel- forest at Palo Verde, Granacaste Province. The results are opment Oetween species occurring in the same area under being prepared for publication. The generalization by varying moisture conditions (snow depth) were evident, Janzen (1967) that flowering in the highly seasonal cli- as were differences between the two areas. Additional mate of Granacaste is concentrated in the dry season is precipitation (snow) delays the emergence and flowering shown to be incomplete. At Palo Verde 35 species flow- of many species (Donald Collins, personal communi- ered in the dry season, 36 in the wet season, and 13 in cation). both seasons. The flowers that appear in the dry season tend to be more conspicuous than those that appear in Phenology and Forest Genetics the wet season. Most trees mature their fruits in the dry Foreft geneticists must understand the relationship season. Correlations are being worked out between the phenology of the woody plants and their pollination sys- between (1) budbreak and late frost in spring and (2) tim- tems, their seed dispersal systems, and the circumstances ing of the development of male and female cones_ Timing of opening of the vegetative bud is one of the most criti- of their seedling establishment. This activity, in which cal factors in the growth of the white spruce seedling_ If H. G. Baker is the principal investigator, is part of an eco- the bud opens too soon, the young, tender needles are system comparison study sponsored by the Organization exposed to late frosts, and the exposure kills the terminal for Tropical Studies with the aid of a grant from the Na- tional Science Foundation. growth. The breeder's objective is to avoid frost damage by selecting fast-gowing clones, which produce late- Root and Shoot Phenology flushing buds. Forest Service investigators in Wisconsin observed flushing and growth of 27 progenies in the Study of the phenology of root growth and leaf de- greenhouse and correlated these results with phenological velopment is providing a better understanding of the re- observations in the field. Field observations had previ- lationship between two major northern hardwood forest ously indicated that in northern Wisconsin frost might tree species: yellow birch (Benda lutea) and sugar maple kill buds as often as three times in one spring and might (Acer saccharum). Sugar maple root activity is at a peak occur in certain planting sites as late as early July. early in spring while leaves are expanding. The beginning Analysis of the phenology of budbreak suggests of root activity in yellow birch may lag behind that of that control of this event is a function of temperature sugar maple by as much as 2 weeks, and peak root activ- and the genetic constitution of the tree. Differences in ity may lag as much as a month. These two species are temperature summation from one spring to another cause competitors, and knowledge of their relationship in the differences of as much as 2 weeks in dates of budbreak. northern hardwood forest is critical in forest manage- Knowing the origin of seeds does not always make it ment. Sugar maple may have a competitive advantage possible to know what the date of flushing will be. Seeds over faster-growing species as a result of early root from the same source often produce trees that differ con- growth (C. Tubbs, personal communication). Phenologi-

:447 cal studies of roots are being continued to relate the be- Studies similar to those in the Lake Wingra Basin have ginning and continuation of root activity to environmen- been initiated at Oak Ridge, Tennessee. Information is tal factors. If a correlation is found, it may be possible to being accumulated to permit the selection of phenologi- predict patterns of root activity without making obser- cal indicator events. Records for the first 7 years have vations. been published (Taylor, 1969). Phenological studies are planned in the Lake George Effect of Fertilization on Phenological Events watershed, another area included in the deciduous forest biome program. The observations of established species A study in the northern Great Plains is relating crop will be keyed to those at Lake Wingra and Oak Ridge, phenoiogy to the application rates of nitrogen on range sites in the northeastern lilac network, and Indiana and sites. Knowledge of the cffects of fertilization on pheno- other midwestern bench-mark stations. logical events would make it possible to use both fertil- In the Texas grasslands, observations are being made izer and native grassland range more efficiently. One on vegetation, reproductive stages, and phenology as re- benefit might be a reduction in frost damage. For exam- lated to climate and soil moisture. Initial work in Nevada ple, if a farmer knew that heavy applications of nitrogen suggests that differences in timing of events in that region to grass would result in the emergence of tender growth are due primarily to site and precipitation and secondar- when frost was expected, he could reduce the likelihood ily to relationships between development rate, soil mois- of frost damage by applying nitrogen at lower rates. ture depletion, and soil temperature. In the cold desert, Long-tenn phenological data may also provide informa- the overriding influence on phenology appears to be the tion by which the quality of range sites can be catego- precipitation-soil moisture factor. Thus, there are re- rized (Goetz, 1969). Adding fertilizer to a native grass- gional differences in the major factors controlling pheno- land can increase utilization by livestock. Many species logical events (Bro. Daniel Lynch and Paul Tueller, per- of plants are present in native grasslands. The timing of sonal communications). phenological development of the various species must be known so that the most desirable plants receive the bene- Relationships between Insect and Plant fit of the fertilizer. There is a related need for a more Development fundamental understanding of the morphology of native grasses and their response to grazing (Green,1969). The studies reported in the preceding sections are concerned with plants and plant development. Pheno- Phenology in Ecosystems logical information is also being obtained on animals, birds, and insects and on relationships between plants Phenological data often may be substituted for other and insects. In Wisconsin the developmental sequencing biological data or for meteorological data. For instance, of several mosquito species was studied in relation to the the deciduous forest biome study in the Lake Wingra development of a variety of plant indicator species. Pre- Basin (IBP) will use phenological data from the Univer- liminary results suggest that the time of appearance of sity of Wisconsin Arboretum and nearby areas to com- certain mosquitoes and other Diptera may be signaled by pare and coordinate seasonally related sequencesof the development of certain plant phases (DeFoliart et al., various ecological processes at different points within 1967). the basin.* Phenological data are easier and cheaper to In Texas the first emergence of Culex Tarsalis females obtain than are meteorological data. The considerable appears correlated with the flowering of spike rush and body of data available for analysis will be supplemented tansey mustard, and the first isolations of encephalitis with information from the Wisconen Phenological So- from sparrows were correlated with the first appearance ciety and from investigations on vegetation and meso- of chimney swifts and with initial flowering of several climate.t plants (e.g., arrowhead, curly dock, and pink smartweed). The coordinated study of phenology is a step toward Preliminary data suggest that there may be worthwhile prediction of primary production in all the biomes in- correlations between important stages in the seasonal his- cluded in theIBPanalysis-of-ecosystems program. Sites tory of arboviruses and phenological phenomena of birds may be related by phenological events, differences in and plants within the same ecosystem (A. D. Hess, per- length of the growing season, and limiting factors, all of sonal communication). which will be monitored. Bird and Animal Phenology *J. A. Zimmerman. 1969_ "A 35-Year Record of Selected Spring Events at Madison, Wisconsin, 1935-1969." Presented at 1969 Other studies are investigating the phenology of ani- meeting of the Wisconsin Academy of Sciences, Arts and Letters. mal development relative to climate. A study of the 'CO. L. Loucks and Nancy Knight. 1968. "The Mesoclimate of movement of blackbird populations in northern Ohio Wisconsin in Relation to Natural Vegetation." Final report of the project in environmental sciences. University of Wisconsin. provides data both on the sequence of movements by

88 these birds and on thning of these movements relative ment, and synagraphic computer mapping. BioScience 21:62-70. to climatic conditions and vegetational development. Nienstaedt, H., and J. P. King. 1969. Breeding for delayed bud- A study on the reproductive phenology of redwing black- break in Picea glauca (Moench) Voss-Potential frost avoid- birds in upland habitats is in progress. It examines the ance and growth gains. Second World Consultation on Forest Tree Breeding. 14 p. effect of microclimate on laying, nest construction, and Taylor, F. G. 1969. Phenological records of vascular plants at nesting development. Results will enable wildlife manage- Oak Ridge, Tennessee. Deciduous Forest Biome, IBP. Oak ment specialists to make predictions about the effect of Ridge National Laboratory. ORNL-IBP-69-1. 46 p. weather on nesting and on the survival of nestlings (M. I. Dyer, personal communication).

APPENDIX: CONTRIBUTORS

Forest Stearns, Department of Botany, University of Wisconsin, Milwaukee (Program Director) REFERENCES Byron 0. Blair, Department of Agronomy, Purdue University, Lafayette, Ind. Addison, E. S., HI. 1969. Forecast of blueberry harvest dates as Joseph M. Caprio, Plant and Soil Science Department, Montana determined by climatology. M.S. thesis, Rutgers University. State University, Bozeman Caprio, J. M. 1966. Patterns of plant development in the western Donald Collins, Department of Botany and Microbiology, Mon- United States. Montana Agr. Exp. Sta. Bull. 607. 42 p. tana State University, Bozeman Caprio, J. M. 1968. Instructions for selecting a common purple Gene R. DeFoliart, Department of Entomology, University of lilac bush for making observations on lilac learmg and bloom Wisconsin, Madison dates. Montana Agr. Exp. Sta. 1 p. M. I. Dyer, Ohio Blackbird Research Laboratory, Bureau of Sport Caprio, J. M., M. D. Magnuson, and H. N. Metcalf. 1970. Instruc- Fisheries and Wildlife, U.S. Department of the Interior, tions for phenological observations of purple common lilac Sandusky, Ohio and red-berry honeysuckle_ Montana Agr. Exp. Sta. Cir. 250. Harold Goetz, Department of Botany, North Dakota State Uni- 19 p. versity, Fargo Caprio, J. M., and H. N. Metcalf. 1970. Report to cooperators in A. K. Helium, Department of Fisheries and Forestry, Calgary, the phenological survey in the western region of the United Alberta, Canada States. Montana Agr. Exp. Sta. 7 p. A. D. Hess, Communicable Disease Center, U.S. Public Health DeFoliart, G. R., M. R. Rao, and C. D. Morris. 1967. Seasonal Service, Fort Collins, Colo. succession in blood sucking Diptera in Wisconsin during 1965. Richard J. Hopp, Department of Plant and Soil Science, Univer- J. Med. EntomoL 4:363-373. sity of Vermont, Burlington Goetz, H. 1969. Growth and development of northern Great R. E. Jones, Department of Entomology and Applied Botany, Plains species in relation to nitrogen fertilization. J. Range University of Delaware, Newark Manage. 23:112-117. Ronald Labisky, Illinois Natural History Survey, Urbana, Ill. Green, C. E. 1969. Morphological variation in three ecotypes of Katherine Lettau, Secretary, Wisconsin Phenological Society, Agropyron Smirhii Rydb. and Bouteloua gracilis (H.B.K.) University of Wisconsin, Madison Lag. Ex. Steud. M.S. thesis, North Dakota State University. Helmut Lieth, Department of Botany, University of North Caro- 83 p. lina, Chapel Hill Helium, A. K. 1967. Periodicity of height growth in white spruce Orie Loucks, Department of Botany, University of Wisconsin, reproduction. Forest. Chron. 43:365-371. Madison Hopp, R. J., M. T. Vittum, and N. L. Canfield. 1969. Instructions Bro. Daniel Lynch, Department of Biology, St. Edward's Uni- for phenological observations: persian lilac. Pamphlet 36- versity, Austin, Tex. Vermont Agr. Exp. Sta. 8 p. Hans Nienstaedt, Institute of Forest Genetics, U.S. Forest Ser- Janzen, D. H. 1967. Synchronization of sexual reproduction of vice, Rhinelander, Wis. trees within the dry season in Central America. Evolution Carl Tubbs, Northern Hardwood Laboratory, U.S. Forest Service, 21:620-637. Marquette, Mich. Lieth, H. 1970_ Phenology in productivity studies, p. 29-46. In Paul Tueller, Renewable Resources Center, University of Nevada, D. E. Reichle [ed.] Ecological studies. VoL 1. Springer-Verlag, Reno Berlin. Walter G. Whitford, Department of Biology, New Mexico State Lieth, H., and J. S. Radford. 1971. Phenology, resource manage- University, Las Cruces

88 Conservation of Plant Genetic Materials

Modern crop husbandry is based on a relatively small In 1969 and 1970 FAO held conferences in Rome to segment of the plant kingdom. These highly efficient develop plans initiated at the 1967 conference. These products of man's research, however, still depend on the conferences resulted in formation of an FAO Crop Ecol- genetic resources that occur in the "gene centers" of the ogy and Genetic Resources Branch with headquarters in world. Gene centers may be either the regions that repre- Rome. This unit coordinates international efforts in germ sent the geographic origin of the species or areas where plasm conservation. primitive varieties have been grown. Many of these pro- In 1968 a symposium on germ plasm was held as a genitor types or weedy relatives of crops have been the part of a meeting of the American Institute of Biological principal sources of pest resistance or other essential Sciences at Columbus, Ohio. About 10 papers were pre- traits. sented on the status of the germ plasm base in the United Gene pools are seriously threatened with destruction, States and on possibilities of conserving the base through by overgrazing and related malpractice, and with loss participation in the IBP. The papers wei published in through replacement of land races" by improved vari- Economic Botany 23:4 (1969). eties. Once the primitive races are destroyed or replaced, In December 1969 the U.S. National Committee for they can never be recovered. the IBP recommended that an integrated research pro- Recognizing that these resources are essential to agri- gram on conservation of plant genetic materials be in- culture in advanced and developing countries alike, inter- cluded in the U.S./IBP. The Director of the Program is national organizations concerned with food production John Creech. encourage the development of national efforts under the International Biological Program (IBP) to attack the problem of dwindling germ plasm reserves in a coordinated fashion. OBJECTIVES Except for sunflower and some minor crops, the United States does not have a natural base of crop germ plasm. Since our nation's beginning, we have had free ac- The objectives are: cess to the world's germ plasm, and we have made greater To conduct national surveys of primitive and wild use of introduced species and varieties than has any other piant genetic resources held in the United States and to nation. Now when there is an urgent need to ensure con- determine the scope, quality, and usefulness of these tinuance of the availability of these resources, we fmd resources that agricultural practices may restrict or obliterate them. To make full use of automatic data processing In September 1967 the Food and Agriculture Organi- techniques in assembling information gained in the sur- zation of the United Nations (FAO) and the IBP spon- veys and in analyzing, storing, and retrieving it sored an international technical conference to determine To disseminate results of the surveys to IBP, FAO, courses of action to conserve crop germ plasm. The con- the Agency for International Development (AID), and ference found that a world survey of threatened germ other organizations plasm should be conducted, that appropriate institutions To cooperate in national programs to use and per- should serve as centers for advanced training in plant ex- petuate our plant genetic resources ploration and collection, and that international programs To cooperate with IBP, FAO, AID, and other or- for storing and retrieving genetic information relating to ganizations in developing priorities for collecting germ germ plasm collections should be encouraged. plasm in threatened biological and geographical areas

90 RESEARCH PLANS held under optimum conditions to ensure long-term viability and are released only when working stocks are unavailable. A reserve storage includes all crops, while This program consists of three activities: surveys of working stocks are usuaily restricted to single crops. collecidons, data handling, and conservation and use of germ plasm.

RESEARCH METHODS AND PROGRESS SURVEYS OF COLLECTIONS

Surveys of collections of genetic stocks in federal, The Agricultural Research Service (ARS), in coopera- state, and private institutions are required as a basis for tion with AID, has developed programs in Puerto Rico assessing the need for exploration and accumulation of and elsewhere for assembling, evaluating, and distributing germ plasm. FAO will integrate the surveys in the United breeding stocks of edible yams and other root crops. States with similar surveys in other countries. Thus, the Similar programs are under way on pulses (edible leg- United States will have inventories of stocks in this umes) and sorghurns. Each program uses existing col- country and will have access to information gathered in lections of seed or vegetative stocks and expands on these other countries. bases through exchange or collection of additional stocks. The information compiled by this program makes pos- The goal is to provide authentic, disease-free breeding sible research on centers of diversity and broadens our materials. understanding of the movement to and establishment of The Rockefeller Foundation, in cooperation with crops in secondary centers of diversity and other aspects ARS and several private institutions, has initiated na- of man's effects on crop dispersal. tional surveys of wheat, corn, sorghum, and rice. In keeping with plans for a global network of breeding research to upgrade crop production in developing countries, the DATA HANDLING foundation has initiated a program to increase pulse germ plasm. To be of maximum usefulness, data concerning col- ARS, in cooperation with FAO, the Rockefeller Foun- lections of genetic stocks must be handled through auto- dation, and other agencies, is assessing the need for ad- matic data processing techniques. The program on conditions to existing wheat collecEions. A statement of pro- servation of plant genetic materials has equipment for cedure has been accepted. The existing wheat collections accumulating information, analyzing it, and preparing lack some varieties from the Near East, and plans are printouts. The printouts will provide information on being developed to collect some of these varieties. variaions in collections, on sources of germ plasm, and The world collections of sorghum are inadequate and on geographic patterns of resistance to diseases and in- incomplete. The most serious deficiencies are lack of wild sects. Information thus made available is an aid in estab- and weedy races, incomplete collections in Ethiopia and lishing priorities for plant exploration and in identifying Sudan, a poor collection of Chinese kaoliangs, gaps in the gaps in our germ plasm base. African collection, few collections from West Pakistan, poor collections from Southeast Asia, poor representa- tion of grassy and fodder types, and lack of collections CONSERVATION AND USE OF of species in other sections of the genus. Uncollected GERM PLASM germ plasm is threatened, but the danger of immediate loss varies from place to place. Special concern has been The purpose of a gene pool program is to provide expressed for West Pakistan and some Arab communities large blocks of germ plasm to breeders. This variety of in the Near East. Significant erosion is expected in the genetic stock may be required to counteract disease out- near future in Sudan, Ethiopia, and parts of West Africa. breaks or to upgrade crop quality and yield in long-term The extensive collection of maize germ plasm resulting breeding programs. from efforts made in the 1950's will be surveyed and Conserved germ plasm is divided into two categories evaluated by a committee of experts engaged in maize re- on the basis of accessibility: (1) working stocks (world search. Attempts will be made to determine the parts of collections) held in adequate storage and available when the world where additional collecting is needed to ensure needed and (2) reserve stocks held in such facilities as the preservation of existing strains and populations. Mainte- National Seed Storage Laboratory, Fort Collins, Colo- nance and distribution of germ plasm will be studied in rado, for long-term preservation. The reserve stocks are an effort to find areas in which improvements are needed.

90 Efforts are under way to bring into existing germ plasm Plant Science Research Division have been assembled by banks the extensive but scattered collections that have the Program Director. The reports show that commodity been made in Europe during the past decade. branches of the Crops Research Division hold an esti- Data on numerous characteristics of 4,829 bean intro- mated 113,000 lines as seed in working collections with ductions have been stored in an automatic data retrieval an additional reserve in the National Seed Storage Labo- system developed at the Taximetrics Laboratory on the ratory of 78,000 accessions. Similar information from campus of the University of Colorado, Boulder. The bean holders of germ plasm in experiment stations was as- records include data on 26 plant characteristics and fam- sembled by J. R. Harlan for FAO. ily, genus, species, and genetic information identifying a species hybrid. They also include information on origin (country and locality) and on the local variety name. Requests for material by characteristic can be processed within a minute. Ar .Avers to the queries appear on a APPENDIX A: PUBLICATIONS printout. The cost of each request to the information Creech, J. L. 1970. Tactics of exploration and collection. In bank is likely to be less than the cost of typing a letter. Genetic resources in plants: their exploration and conservation. IBP Handbook No. 11. London. The longitude and latitude of the collection sites for Harlan, J. R. 1969. Ethiopia: a center of diversity. Econ. Bot. most of the Phascolus vulgaris accessions from Turkey 23(4):309-314. have been determined. A plotter was used, with a com- Hyland, H. L. 1970. Description and evaluation of wild and primitive introduced plants. In Genetic resources in plants: puter map of Turkey, to show the geographic distribution their exploration and conservation. IBP Handbook No. 11. of the sites. A computer program is being worked out London. whereby beans having a particular characteristic or com- Knowles, P. F. 1969. Centers of plant diversity and conservation of crop germ plasm: safflower. Econ. Bot. 23(4):324-330. bination of characteristics can be plotted as to origin. Konzak, C. F., and S. M. Dietz. 1969. Documentation for the This type of study can be applied to records on collec- conservation, management, and use of plant genetic re- tions from any part of the world. sources. Econ. Bot. 23(4):299-309. Reitz, L. P, and J. C. Craddock. 1969. Diversity of germ plasm The data on the beans from Turkey are especially sig- in small Fain cereals. Econ. Bot. 23(4):315-324. nificant even though most of the stocks were collected at Rowe, P. R. 1969. Nature, distribution, and use of diversity in bazaars. Dr. and Mrs. Basri Devecioglu, of Izmir, Turkey, the tuber-bearing Solanum species. Econ. Bot. 23(4):330-.339. who located the sites, stated that a bazaar could be expected to have beans grown primarily in a nearby area. As the information on disease reaction and other aspects of the collection is accumulated, this work may serve as a basis for a better understanding of the ecological and APPENDIX B: PARTICIPANTS sociological factors that affect the culture of plant John L. Creech, Plant Science Research Division, Agricultural varieties. Research Service, U.S. Department of Agriculture, Beltsville, The Rockefeller Foundation and ARS are cooperating Md. (Program Director; germ plasm collections; exploration) William R. Brown, Pioneer Hi-Bred Corn Company, Des Moines, in a pulse increase program, using working collections of Iowa (corn; working collections; surveys) beans, pigeon peas, chick peas, mung beans, and other T. T. Chang, International Rice Research Institute, Los Balms, types that are common in the diets of developing coun- Laguna, Philippines (rice; working collections; surveys) Jack R. Harlan, Department of Agronomy, University of Illi- tries. The increase is designed to build up working stocks nois, Urbana (sorghum; working collections; surveys) for preliminary evaluation and subsequent use in breed- Russell T. Johnson, Spreckels Sugar Company, San Francisco, ing programs. More than 20,000 collections are involved. Calif. (germ plasm survey coordination) Calvin Konzak, Department of Agronomy, Washington State Work is being done at the Federal Experiment Station, University, Pullman (data banks) Puerto Rico, and in Colombia and Nigeria. Louis Reitz, Plant Science Research Division, Agricultural Re- search Service, U.S. Department of AFiculture, Beltsville, The Crop Ecology and Genetic Resources Branch, Md. (wheat; working collections; surveys) FAO, has initiated an international survey of holders of Ralph Richardson, Jr., Rockefeller Foundation, New York, N.Y. crop germ plasm. In the United States, reports for the (pulses; root crops; general germ plasm coordination)

92- Conservation of Ecosystems

Along with increased public interest in environmental aquatic, for scientific purposes. The program will include quality has come a desire to set aside and protect samples disturbed areas, natural areas, and areas that have a long of natural environments. The major thrust has been di- history of scientific study. rected toward preserving areas that have aesthetic or rec- There are three immediate goals: reational value. Interest in areas having scientific value To prepare an inventory of environments that have has been secondary. been set aside for research purposes and to identify other Individua; citizens, groups of citizens, and government areas that should be set aside in order to ensure the pro- agencies (federal, state, and local) have devoted a great tection of a complete, representative series of major eco- deal of effort to acquiring natural areas, but they have systems often depended on inadequate criteria in determining the To develop management practices for preserving quality of the areas. They sometimes have had little idea areas that have been set aside of the value of areas already acquired, and often have not To establish a clearinghouse procedure by which developed sound plans for managing the areas. information concerning research natural areas in the No one really knows how many natural areas have United States can be made readily available to scientists been set aside in the United States, what kinds of areas they are, what additional kinds of areas should be set RESEARCH METHODS AND PROGRESS aside in order to have a complete series of environmental systems, or what kinds of areas are still available for pres- FEDERAL LANDS ervation. The great need is for more effective coordination and greater insistence on applying sound criteria to A Federal Committee on Research Natural Areas rep- evaluate the research values of the areas. resenting the Forest Service, the Bureau of Land Manage- Areas preserved for long-term scientific use provide ment, the National Park Service, the Bureau of Sport natural outdoor laboratories for analyzing many kinds of Fisheries and Wildlife, the Department of Defense, and ecosystems. The environment in these natural or semi- the Atomic Energy Commission was organized early in natural areas can be compared with more disturbed en- 1966 to inventory certain protected natural areas on fed- vironments and thus can serve as a means of measuring eral landsthose having special research valuesand to changes in environmental quality. The areas also provide compile a directory of these areas. The federal govern- opportunities for conducting research on the use, pro- ment holds title to slightly more than one third of the tection, and management of comparable terrestrial or land in th. ilted States, most of its holdings being in aquatic areas. Data thus obtained are valuable as a basis the West and in Alaska. Federal lands are administered by for guides to resource management, such as lf.:nd capa- a number of agencies for a variety of purposes. Most of bility classifications. Many such areas are the source of the biotic, geologic, pedologic, and aquatic types of this animals and plants used in medical or agricultural experi- country occur on these lands and present almost unlim- ments. (See "Conservation of Plant Genetic Materials," ited potential for natural science research. p. 89.) Finally, these areas may be the last protected The Committee was unable to find a system of classi- habitats of endangered animal or plant species. fication that satisfactorily characterized the scientific The conservation-of-ecosystems program is concerned features that the natural areas had been established to with all kinds of ecosystems. In some ecosystems, man is preserve. Thus, it became necessary for the Committee the dominant environmental factor; in others, he is less to accept a composite of several systems. important; in still others, he is essentially an intruder. When the initial inventory was begun, few of the The Program Director is G. Sprugel, Jr. agencies had progressed far in identifying and setting aside research natural areas for special treatment. Utiliz- OBJECTIVES ing existing data, the Committee published A Directory of Research Natural Areas on Federal Lands of the United The chief objective of this program is to describe and States of America in the fall of 1968. The Directory preserve a variety of environments, both terrestrial and (available from the U.S. Government Printing Office)

92 identifies, classifies, and briefly describes 336 areas rep- MARINE AND ESTUARINE AREAS resenting examples of forest, range, and aquatic communities and geological features that had been set aside for J. E. Randall, Bernice P. Bishop Museum, Honolulu, scientific and educational purposes. is responsible for compiling a catalog of marine and estu- The Committee has been reorganized and enlarged arine areas of scientific interest. He has begun to develop under the leadership of R. E. Buckman, U.S. Forest Ser- criteria for selecting the areas and to bring together the vice, and has expanded its efforts to identify additional available information on U.S. marine sites.* Seventy-six research natural areas worthy of preservation. marine sanctuaries established under state laws have been An effort is being made to locate and set aside some identified in 13 states and 1 territory. Eighty-seven more 325 additional areas to ensure the protection of ecosys- have been proposed, and others are being studied. An ad- tems inadequately represented among the existing pre- ditional 17 marine or estuarine sanctuaries have been served sites. proposed in states and territories where none have been Checksheets will be used in gathering more detailed established to date. Seventeen federal marine sanctuaries information on the federal areas. Information thus ob- have been identified. tained will be more comprehensive than that given in the Directory. WORLDWIDE INVENTORY

NONFEDERAL LANDS A worldwide inventory of natural preserves is being compiled under the supervision of the Conservation Ter- The most difficult part of the conservation-of- restrial Section of the International Biological Program. ecosystems program is the inventory of terrestrial re- The objective is to describe the ecological characteristics search natural areas on nonfederal public lands and on and the special biological or conservation features of privately owned lands. During the summer of 1968, the each area. Checksheets are used to collect the information. Conservation Foundation assisted in this task by em- Checksheets have been distributed to federal agencies ploying M. J. Pontrelli, University of Nevada, to assess and other responsible organizations for completion. The the scope of the undertaking and to identify sources of agencies are providing information on research natural information. Dr. Pontrelli prepared a report that gives, areas and on some wilderness areas that are included in for many states, some sources from which information the inventory of U.S. federal lands (see "Federal Lands," on nonfederal terrestrial research natural areas can be ob- above). About 200 checksheets have been completed and tained. The report reveals that there is no central source forwarded to IBP's Monk's Wood Data Center. of such information in 27 states. However, much of the needed information can probably be obtained from organizations and individuals in these states. Paul Lemon, formerly of the State University of New York at Albany, has been employed to complete the task APPENDIX: PARTICIPANTS with funds that became available in the fall of 1970. G. Sprugel, Jr., Chief, Illinois Natural History Survey, Natural Resources Building, Urbana (Prop-am Director) R. E. Buckman, Forest Service, U.S. Department of Agriculture, Washington, D.C. (Deputy Program Director; Chairman, AQUATIC AREAS Federal Committee on Research Natural Areas) R. M. Darnell, Department of Oceanography, Texas A & M Uni- versity, College Station In 1966, R. M. Darnell, Department of Oceanography, F. R. Fosberg, Special Advisor on Tropical Biology, Smithsonian Texas A & M University, undertook the task of devel- Institution, Washington, D.C. oping a program for identifying, describing, and cata- Paul Lemon, American Institute of Biological Sciences, Wash- ington, D.C. loging freshwater areas in the United States that have sci- G. E. Likens, Division of Biological Sciences, Cornell Univer- entific value. Plans for conducting a survey of freshwater sity, Ithaca, N.Y. sites were made, and scientists in various parts of the J. E. Randall, Bernice P. Bishop Museum, Honolulu, Hawaii R. S. Sigafoos, Research Botanist, Geological Survey, U.S. country were invited to participate. G. E. Likens, Cornell Department el' the Interior, Washington, D.C. University, headed a team that made a detailed survey of L. M. Talbot, Council on Environmental Quality, Washington, freshwater sites in the Northeast. Some 75 freshwater D.C. sites in 22 states have already been recommended to the IBPfor international recognition as aquatic natural areas. *J. E. Randall. 1969. Conservation in the sea; a survey of marine Checksheets have been completed on four sites. areas. Oryx 10:1.

93 HUMAN ADAPTABILITY COMPONENT Biosocial Adaptation of Migrant and Urban Populations

The importance of studying migrants from rural to ization is increasing. Peoples who have not shared fully urban areas is emphasized by the extraordinary changes in such movements in the pastAmerican Negroes, Eski- in the distribution of populations throughout the world. mos, Yugoslavian mountaineers, Russian peasants, These changes are being forced by rapid social, political, Chinese villagersare being swept into an industrialized, and economic developments. Under the pressure of new impersonal society with little preparation for such a dif- technologies and rising expectations, the pace of change ferent life. On the whole, the economic rewards have is accelerating, and rural or isolated peoples are trans- been considerable, but the social costs have been great. formed ovemight into urban dwellers. Primitive tribes are Important matters of public policy and of scientific con- brought into shattering contact with modem civilization. cern are involved. Should the process of urbanization be Megalopolis expands at the expense of the country- accelerated or retarded? How can the stresses of assimila- side. The growth of suburbs has its counterpart in the tion in new communities be reduced? What is the effect desertion of rural villages and the return of farmland to upon a village of losing its youth? Does the city rob the woods. Shifts in population, no less notable in the devel- countryside of the most innovative? What are the limits oping than in the industrialized countries, are altering es- of human ability to adapt to new environments, and to tablished ecological balances and subjecting increasingly what degree are these determined by genetic makeup, by larger groups to the stresses of migration and assimilation. biological constitution, by psychological set, by age, and Important migrations are occurring throughout the by education? These are some of the questions with world, but they have been subjected to little sophisti- which this program is concerned. Others are posed in the cated analysis. In particular, the biological and psycho- statement of objectives that follows. logical aspects have been ignored. Many writers have re- The Program Director is Everett S. Lee. marked upon the success of the overseas Chinese, the overseas Indians, and the Japanese-Americans. In contrast is the distress of blacks in metropolitan ghettoes and of southem whites in Cleveland and Cincinnati. In Latin OBJECTIVES America and Asia, villages lose many of their youth to the barriados, or urban slums. Migration causes dramatic changes in the composition The objectives of this program are to fmd the answers of local populations. There are counties in the United to the following questions: States where more than 70 percent of the persons who How do migrants differ from nonrnigrants in the were 10-14 years of age in 1950 had left by 1960; many area of origin? From nonmigrants in the area of destina- left as soon as they finished high school or dropped out. tion? From migrants in other migration streams? From In some counties there are more people aged 70-79 than migrants of earlier times? there are aged 20-29, and in at least 300 counties there What is the impact of migration on the structure of have been years in 1.1,aich there have been more deaths the population at origin? On the structure of the popula- than births. The United States is not the only country in tion at destnation? On the social and psychological mi- which the cities grow at the expense of the rural areas. In lieu at origin and destination? Argentina, for example, there are villages from which What is the impact of migration on the psychologi- more than half of the youths migrate. cal and physical state of the migrants? On the psychologi- There has been a vast, worldwide movement of popu- cal and physical state of the migrants' children? lations from rural to urban areas, and the rate of urban- What changes in social and economic conditions

9)5 have the greatest effect on the volume of migration? On dwelling area, religion, and social class will be completed the selection of prospective migrants? On the types of for each available nonwhite family. Random samples migrants' origins and destinations? drawn from this universe will be asked to participate in What are the most important factors in the assimi- a health survey that will include height,weight, blood lation of migrants at destination? pressure, chest X ray, electrocardiogram, visualacuity, What are the most important obstacles to the as- gross physical impairments, fat pad thickness, breast ex- similation of migrants? amination, hemoglobin, hematocrit, tests for sickle cell, and indications of malnutrition. It is expected that about 750 migrants will be followed by the University of Illinois College of Medicine. Success or failure in Chicago will be assessed. It is hy- RESEARCH PLANS pothesized that successful migration and adjustment in Chicago will be associated with: A hill rather than a delta origin (nonplantation Research is planned that will highlight differences in rather than plantation) response to environmental changes in different cultures Above-average education of parents and clarify differences in mobility among different types Active religious participation by the family of people in the same culture. Special emphasis is placed Presence of both parents in the home until all off- on studies of groups in extreme or rapidlychanging en- spring are at least 9 years of age vironmentsfor example, the Eskimo, the rain forest Nonpossession of a criminal record by parents native, or the peasant in a society that is industrializing. Migration as a family unit The selection of migrants at origin and their assimilation Successful relatives in Chicago at destination will be the focus of several studies. Migrants in Chicago will be followed over a 3-year Special attention will be paid to biological or health period, and nonmigrants in Holmes County will be fol- factors. It is well known that social and psychological lowed over several years. factors are important in migration and urbanization, but This project is unusual in that the cooperation of the little is known about physiological state or intellectual Holmes County residents has been obtained, and some capacity as determinants of migration. Experiments will Holmes County residents will participate in the investi- be made to see if genetic markers can be used to differen- gation. The study is directed by anthropologists and tiate migrant from nonmigrant, and residual populations medical people from the University of Illinois. in areas of high out-migration will be examined for possible high incidences of physical or mental defects, some of which may be genetic in origin. Migrants from Yazoo County This study is concerned with the characteristics of migrants from Yazoo County, Mississippi, to Chicago and other cities. It is financed by the Office of Economic RESEARCH METHODS AND PROGRESS Opportunity. The reasons for migration and assimilation difficulties at destination are explored. Migrants are located and interviewed at destination, and special empha- STUDIES UNDER WAY sis is placed on nutrition and dietary habits. This study and the Holmes County study give a picture of the black The following studies are under way: migrants from migrant from the rural South in northern cities, a picture Holmes County, Mississippi; migrants from Yazoo County, more detailed than any previously obtained. Such studies Mississippi; and adjustment of peasants to industrial life are important because of the continuing exodus of young (Yugoslavia). people from rural areas in the South to large cities outside the region. Contact with large cities changes the outlook in the Migrants from Holmes County communities of origin. In Yazoo County, as in Holmes This study was originally designed in part as a segment County, numerous migrants have returned from Chicago, of the IBP's study of migrant peoples. Data from a com- and a high proportion of the black population has visited plete sample of the black population of Holmes County Chicago or has had visits from relatives who migrated will iv! studied to assess the effects of migration on the there. Initial findings from both studies suggest that mi- health of rural Negroes who move from Holmes County grants to Chicago from these poor Mississippi counties to Chicago. Detailed questionnaires on family size and are more successful and less likely to be on welfarethan structure, educadon, occupation, income, type of house, are blacks raised in Chicago ghettoes.

95 Adjustment of Peasants to Industrial Life An industrial medicine team remains in Ivangrad throughout the year and collects information on each This study is sponsored by the Federal Institute of worker's ability to meet quotas and get along with other Public Health, Yugoslavia, and by the National Center workers, his supervisors, and townsmen. The records for Health Statistics, U.S. Public Health Service. It was maintained by the factory management are made avail- initiated by Mitomir Savieevie and Momeilo Kilibarda able for each worker. (see Appendix, p. 97). The locale of the study is Montenegro, where there is an unusually good opportunity to study the adjustment FUTURE STUDIES of peasants to city life and industrial pursuits. In the last few years the one-time Turkish garrison town of Berena The following studies will be proposed for funding: has been transformed into the industrial city of Ivangrad, cohort of 1948 (Sweden), assimilation of immigrants where a large paper and cellulose mill, a plywood and from different countries and the adjustment of their chil- plywood products factory, a leather factory, and a c..)al dren (Israel), assimilation of native peoples in cities mine furnish employment for about 2,000 workers, most (Alaska), and migration of Appalachian whites to mid- of whom were drawn directly from peasant life. The western cities_ transition in many cases was abrupt, from sheep herding one day to work in the factory the next, from life in a Cohort of 1948 mountain hut to an apartment in a modern high-rise building, from providing nearly all of one's food to buy- This study is planned as a continuation of Thorsten ing food in a supermarket. Husen and Ejnun Neymark's study of migrants and non- All of the 2,000 factory workers have been given a migrants in Sweden. Their study was concerned with the series of psychological, physical, and mental examina- entire cohort of males born in Sweden in 1928 and called tions, and all have been given 2-hour interviews in their up for possible conscription in 1948. homes. The interview was structured to give a complete Records on a 10 percent sample of the 1948 cohort picture of the worker's childhood and family background, have been kept. They include records from place of birth, together with many items designed to show attitudes to- records from schools, and employment records extending ward country and city life and feelings of pessimism and to 1956. In the IBP study, each of the men in the 1948 optimism. Religion of parents, income, housing, smoking cohort who is still living will be located in the population habits, drinking habits, amusement patterns, friends, registers for 1970. The Swedish system of maintaining family relations, and other items of demographic or so- population registers makes it possible to trace each living cioeconomic importance were also included. person who is still in Sweden and obtain much informa- Psychologists gave an intelligence test, the Cornell tion about him. Medical Inventory, and the Maudsley Personality Test to Reports, including a book published by Neymark, each worker. These tests were followed by a complete indicated that intelligence is a highly significant factor in physical examination and a psychiatric examination. migration and that the higher the intelligence level the A full psychological, medical, and psychiatric report was greater the rate of migration and the longer the distance filed for each person. of migration. Rural communities lose about 90 percent The same interviews and examinations were given of their more intelligent males and retain a high propor- to an equal number of controls selected in the villages tion of the less intelligent. Furthermore, the less intelli- from which the workers had migrated. Testing and ex- gent tend to migrate from large cities to smaller cities or aminations for both workers and controls were first ad- from small towns to rural areas. Similar conclusions hold ministered in 1966, and the procedure was repeated ex- for socioeconomic background. It is evident that migrants actly in 1967 and 1968 for all the workers. At the end have been far raore successful than nonmigrants. of 5 years the workers and the control group will be ex- The work now planned would include following the amined again. Changes in physical and mental condition 1948 cohort to 1970 and would stress the health of mi- will b.! noted. grants and nonmigrants. The differentials that were This is the largest study of its kind ever conducted shown to exist in 1956 will be reexamined. Many health and perhaps the best designed. Nearly all the persons in data are available that have never been fully explored, the sample were interviewed and tested. A monograph and migration will be related to several indices of physi- on the first results of the study is being prepared. These cal and mental health. results indicate a high degree of selection at origin for the Assimilation of Immigrants from most intelligent, the healthiest, and the most optimistic Different Countries among the persons electing factory work. They also have less indication of marked or incipient mental disorder. Population registers in Israel permit following persons'

96 activities in much the same way as in Sweden. In addition returned to their mountain homes will be sought. there are important records that were filed in connection An effort will be made to coordinate the several pro- with immigration and naturalization. Jews from different posals and to see that the materials already collected are countries have had very different degrees of success in quickly exploited. assimilating in Israel, and the children of some groups, particularly those from Asia and North Africa, have had Migrants to and from Aberdeen, Scotland some of the same difficulties in school and in regard to This study will use the records maintained by intelligence testing that have been experienced by Ameri- Raymond Illsley and the faculty of the Medical School, can blacks. The Central Bureau of Statistics has proposed University of Aberdeen. Records have been maintained a study of the assimilation of migrants in which a number on everyone hospitalized in Aberdeen during the last two of socioeconomic indicators would be used to show the decades. Records are also maintained on all persons born degree of assimilation. Fertility and health records of in Aberdeen hospitals. They have created a rester that various kinds would also be considered. School records permits the comparison of migrants with the nonmigrant would be used to study the adjustment of children, and population. Persons migrating to Aberdeen can be com- children raised in the kibbutzim would be compared with pared with those who left Aberdeen. The population in those who live with non-European Jews in the usual the register is now over 20,000. These data show that family situations. migrant women are larger and heavier, have higher IQ's, and have less difficulty in childbirth than nonmigrant Assimilation of Native Peoples in Cities women, and the children of migrant women do better in Alaska is still in a frontier stage, and the effects of in- school than the children of nonmigrant women. truding western culture into native cultures can be ex- The work now planned would extend the present amined in areas undergoing rapid change. comparisons of migrants and nonmigrants and introduce The Institute of Social, Economic, and Governmental better controls for social status and length of residence Research, University of Alaska, proposes a study of the in Aberdeen. assimilation of migrants from Eskimo and other native villages to Anchorage or Fairbanks. Native villages have long been under stress as a result of large numbers of young people abandoning traditional pursuits and taking various kinds ofjobs in industry or entering the armed APPENDIX: PARTICIPANTS forces. A considerable number of girls have learned secre- Everett S. Lee, Department of Sociology, University of Georgia, Athens (Program Director; Yugoslavian study) tarial and clerical skills and have migrated to,the larger James Brown, Department of Sociology, University of Ken- towns or cities. At the same time, infant mortality has tucky, Lexington (Appalachian study) decreased sharply, so that the villages have high propor- Charles Cannell, Field Director, Michigan Survey Center, Uni- versity of Michipn, Ann Arbor (Yugoslavian study) tions of children and older people. Thorsten Husen, University of Stockholm, Stockholm, Sweden It is intended to locate the migrants from the villages (Swedish study) and interview a large sample. School records of the mi- Raymond Insley, Department of Sociology, University of Aber- deen, Aberdeen, Scotland (Aberdeen study) grants will be compared with those of their children. Momalo Kilibarda, Chief, Section of Psychiatry, Federal Insti- Anthropologists will make studies of villages that have tute of Public Health, Belgrade, Yugoslavia (Yugoslavian been most affected by out-migration. Special attention study) Henry Lorenzi, Milton-Olive III Memorial Program for Children, will be paid to villages that are likely to be affected by Lexington, Miss. (Holmes County study) drilling for oil. Bernice Montgomery, Milton-Olive III Memorial Progam for Children, Lexington, Miss. (Holmes County study) Ejnun Neymark, University of Stockholm, Stockholm, Sweden Migration of Appalachian Whites (Swedish study) Daniel O. Price, Department of Sociology, University of Texas, A number of scholars have proposed studies of the Austin (Yazoo County study) migration of Appalachian whites to midwestern cities. 0. K. Sagen, Assistant Director, National Center for Health The University of Kentucky has files that trace the mi- Statistics, U.S. Public Health Service, Washington, D.C. (Yugoslavian study) grants from some communities over a period of several Mitomir Savi6evid, Director, Institute of Industrial Medicine, years. It is evident that migrants to the cities can be lo- University of Belgrade, Belgrade, Yugoslavia (Yugoslavian cated and interviewed, and that they can be compared study) with the nonmigrants in regard to many characteristics. Demitri B. Shimkin, Professor of Anthropology and Geography, University of Illinois, Urbana (Holmes County study) Reasons for migrating or not migrating can be obtained. Moshe Sichron, Central Bureau of Statistics, Jerusalem, Israel Information conceming those who went to the cities but (Israeli study)

97 Nutritional Adaptation to the Environment

Nutritional studies in the U.S. program are integrated is found in a worldwide trend of migration from rural on a national and international basis with other studies areas into congested urban areas where cultural patterns of human adaptation. The results will assist mankind in are confusing and directions of improvements are not making adjustments to present and future circumstances well established. in relation to food. The records of anthropology and history and the contemporary world scene all show clearly that man's relative success in obtaining and consuming food that permits a close approach to optimum health is OBJECTIVE a major factor in efforts to survive and to advance socially and economically. Experience in arctic, tropical, and temperate areas has The objective of this program is to guide mankind to- resulted in three extreme ranges of food practices that ward successful adaptation to environmental changes that are in sharp contrast in regard to critical margins of suc- relate to food. cess or failure in human adaptability. In circumpolar areas, animals often are the only source of food. Because of this restriction, diets are monotonous and close to the RESEARCH PLANS limit of physiological tolerance in their high content of protein and fat and their low content of vitamin C. In tropical areas the reverse is true. Diets there tend to be The program has three subprograms, each of which is derived directly from plant sources; they tend to be ex- related to an area of the world: tropical and subtropical tremely low in good-quality protein and excessively high ecology, temperate ecology, and arctic ecology. Each sub- in starch or sugar; they are often low in total energy, program is coordinated with human population studies minerals, and vitamin A. Temperate areas usually permit that are in progress in each of the areas. more varied and more abunda it food production and The program also includes projects designed to sharpen generally a more favorable economic environment. the tools with which practical corrective measures can be A major penalty on public health in all climates results evaluateu, guided, and accelerated. from excessive caloric intake, which begins in early child- C. Glen King, Institute of Nutrition Sciences, Colum- hood and increases with advancing age. Tne problem of bia University Medical Center, New York, New York, is excessive caloric intake is especially severe in more eco- Program Director. 0. Lee Kline, American Institute of nomically advanced countries_ The regulating factors in- Nutrition, Bethesda, Maryland, is Coordinator. volved should be identified very early in life so that a Support for the management of this program is pro- basis for effective adaptation and health conservation can vided by the Agricultural Research Service, U.S. Depart- be established. Temperate areas also include populations ment of Agriculture. that contend with stresses of altitude, semidesert conditions, migration, and marginal soil deficiencies. Hence, surveys in addition to basic research are an important feature of many international studies. RESEARCH METHODS AND PROGRESS Research to establish the nature and practical limits of human adaptability and to improve food resources accordingly will require recognition of genetic variability TROPICAL AND SUBTROPICAL ECOLOGY and carefully controlled studies with experimental animals to supplement research with human subjects. Developing countries in tropical climates are subject A special feature of nutritional stress and adaptation to a lack of protein that is expressed in the frequent oc-

98 currence of the disease kwashiorkor. Protein calorie mal- Development of Soy-Based Foods of nutrition affects a large segment of the populations in High Nutritive Value the tropics. It is a serious problem in Latin America and Soybeans are an important source of protein. Careful in large sections of Africa, the Middle East, Southeast processing is needed to obtain a product acceptable for Asia, and Pakistan. human consumption. The following projects contribute to our knowledge Soybeans are normally prepared by soaking in water of ways in which the nutritional status of people in the before the grinding operation. The soaking continues tropics can be improved by the identification and treat- over a period of several hours. Enzyme activity during ment of protein deficiency, nutritional anemia and in- this period results in products that cause off-flavors to fections related to undernutrition. develop. One product is 1-octen-3-ol, which has a mush- roomlike odor. This product is reduced by use of water Development of Food from Cellulosic Wastes temperatures over 50° C, but yields of solids in the resulting milks are lowered. Extended studies on the flavor A pilot plant has been constructed in which cellulosic formation and the enzymic causes of flavors have Ied to wastes are converted to microbial proteins. This process a number of suggestions for improved processing. includes a delignification step in which a mild caustic is The rancid flavor of soymilk as traditionally prepared used. The fermentation that follows uses a mixed culture resides in the volatile fraction. Wet-grinding the soaked of cellulomonas and alcaligenes. Cellulomonas is a species beans at 80° C or higher prevents rancidity and results in of cellulolytic organism not previously isolated. a bland-tasting milk low in beany flavor. Preliminary studies indicate that this material is non- The effectiveness of grinding procedures was deter- toxic when fed to rats at a level of 40 rrncent of the diet. mined by particle-size measurements consisting of a com- They also indicate that the product, which contains 70 bination of wet-sieving and sedimentation tests. Hammer percent of protein by amino acid analysis, supports a mills were as effective as attrition mills, and colloid mills good rate of growth in rats. Methionine is the limiting were more effective. Colloid mills reduced the particle amino acid. size to less than 0.05 micron. Research continues to simplify the process and lower The protein redispersibility of dried soymilks was in- production costs. Mutations of the organism are being creased by processing under conditions that split disulfide developed to provide more balanced amino acid compo- bonds and prevented sulthydryl and hydrophobic inter- sition of the protein. actions. The protein will be assessed for acceptability as human Nuts and legumes are roasted primarily to give them a food and for feed and food uses. desirable flavor and to change their texture. In the case of soybeans, roasting also improves nutritional quality by Investigators: C. D. Callihan, W. H. Daly, Y. W. Han, destroying antitrypsin. C. E. Dunlap, V. R. Srinivasan, and W. F. McKnight, Louisiana State University, Baton Rouge, La. 70803 Investigators: W. B. Robinson, L. R. Hackler, J. C. Moyer, K. H. Steinkraus, and J. P. Van Buren, Department of The following publications resulted from this work: Food Science and Technology, New York State Agri- cultural Experiment Station, Cornell University, Callihan, C. D. 1970. How engineers are putting microbes Geneva, N.Y. 14456. to work. Chem. Eng. 77:160-164_ M. C. Bourne, College of Agriculture, University of the Callihan, C. D., and C. E. Dunlap. 1969. The economics Philippines, Laguna, Philippines of microbial proteins produced from cellulosic wastes. Compost Sci. 10:6-12. Han, Y. W., C. E. Dunlap, and C. D. Callihan. 1971. Effect of Undernutrition on Nucleic Acid and Single cell proteins from cellulosic wastes. Food Protein Content of Developing Brain Technol. 25(2):32-35. Han, Y. W, H. A. Schuyten, and C. D. Callihan. 1971. Since the DNA content of diploid cells is constant, Combined effect of heat and alkali in sterilizing sugar total organ DNA reflects the number of cells in that cane bagasse. J. Food Sci. 36(2):335-338. organ. Cell number is calculated by dividing total organ Han, Y. W., and V. R. Srinivasan. 1969. Purification and DNA by DNA per cell. The weight or protein content of characterization of B-Glucosidase from Alcaligenes. each cell is determined by dividing the total organ weight J. Bacteriol. 100:1355-1370. by the number of cells. Srinivasan, V. R., and Y. W. Han. 1968. Isolation and This chemical approach was used in measuring the characterization of cellulose-utilizing bacterium. Appl. pattern of cellular growth in 31 human brains. These Microbiol. 16:1140-1145. were brains of infants and of fetuses having a gestational

100 age of more than 13 weeks. Weight, protein, and RNA Karp, R., J. Brasel, and M. Winick. 1971. Compensatory content increased linearly with time during these periods. kidney growth after uninephrectomy in adult and in- Increase in DNA content leveled off at time of birth and fant rats. Arn. J. Dis. Child. 121:186. reached a maximum at 8 to 10 months of age. This Rosso, P., J. Hormazbal, and M. Winick. 1970. Changes clearly indicates that cell division in the human brain in brain weight, cholesterol, phospholipid and DNA continues to 8 to 10 months of age. Further growth is by content in marasmic children. Amer. J. Clin. Nutr. increase in protein, RNA, and lipid content of each cell. 23:1'775. Following this the regional growth patterns in the Winick, M. 1969. The effect of nutrition on cellular human brain were determined. The data demonstrate growth. In G. Blix [ed.] Symposia of the Swedish Nu- that cell division is most rapid in the cerebrum postna- trition Foundation. VII. Nutrition in preschool and tally. Cell division stops in the cerebrum, cerebellum, and school age. Almquist and Wiksell, Uppsala. brain stem at the same time (8 to 10 months). Winick, M. 1969. Food, time and cellular growth of the In nine infants who died of severe malnutrition during brain. N.Y. State J. Med. 69:302-304. the first year aide, there was a proportional reduction Winick, M. 1969. Malnutrition and brain development. in weight and in protein, RNA, and DNA content of the J. Pediat. 74:667. brain. The number of cells was reduced, but weight per Winick, M. 1969. Nutrition and the ultimate cellular cell and protein content per cell were unchanged. makeup of various tissues. Nutr. News 40:7. In a second group of marasmic children, it was dem- Winick, M. 1970. Cellular growth in intrauterine malnu- onstrated that the reduction in cell number occurred in trition. Pediat. Clin. N. Amer. 17:69. all regions studied and that the most marked effects oc- Winick, M. 1970. Cellular growth of the fetus and pla- curred in the cerebrum. In another study, lipid deposi- centa. In H. Waisman and G. Kerr [ed.] Fetal growth tion was studied. Total cholesterol and phospholipid con- and development. McGraw-Hill, Inc., New York. tent in the brains of marasmic children was also found to Winick, M. 1970. Fetal malnutrition. Clin. Obstet. be decreased. Gynecol. 13:526. These human studies reinforce a series of comparable Winick, M. 1970. Fetal malnutrition and growth processes. studies that have been made on laboratory animals. Hosp. Pract. 5:33. This demonstration that severe early malnutrition re- Winick, M. 1970. Nutrition and mental development. tards cell division and myelin synthesis in the human Med. Clin. N. Amer. 54:1413. brain supports the growing concern over the possibility Winick, M. 1970. Nutrition and nerve cell growth. Fed. that malnutrition in early life retards mental development. Proc. 29:1510. Winick, M. 1970. Nutrition, growth, and mental develop- Investigators: Myron Wmick and Irving Fish, Department ment: biological correlations. Amer. J. Dis. Child. of Pediatrics, Cornell Medical College, 1300 York 120:416. Avenue, New York, N.Y. 10021 Wmick, M. 1971. Cellular changes during placental and Julio Meneghello and Pedro Rosso, Department of Pedi- fetal growth. Amer. J. Obstet. Gynecol. 109:166. atrics, Hospital Roberto del Rio, Santiago, Chile Winick, M. In press. Cellular growth during normal and abnormal development of the brain. In W. Himwich The following publications resulted from this work: [ed.] Biochemistry of the developing brain. Marcel Dekker, Inc., New York. Brasel, J. A., R. A. Ehrenkranz, and M. Winick. 1970. Winick, M. In press. Nutrition and cell growth. In S. A. DNA polymerase activity in rat brain during ontogeny. Miller and M. Winick [ed.] Advances in biochemical Develop. Biol. 23:424. nutrition. Interscience Publishers, New York. Brasel, J. A., and M. Winick. 1970. Differential cellular Wmick, M., and J. A. Brasel. 1970. A "final common growth in organs of hypothyroid rats. Growth 34:197. pathway" for producing either permanent or revers- Duckett, S, R. Rendon, and M. Winick. In press. The ef- ible alterations in organ growth. In J. Masek, K. fects of malnutrition on the developing brain. In P. Osancova, and D. P. Cuthbertson [ed.] Proc. Eighth Black [ed.] Brain damage in children: etiology, diag- Int. Congr. Nutr., Prague, Sept. 1969. Excerpta nosis, management. The Williams and Wilkins Com- Medica, Amsterdam. pany, Baltimore. Wmick, M. and P. Rosso. 1969. The effect of severe Fish, 1., and M. Wmick. 1969. Cellular growth in various early malnutrition on cellular growth of human brain. regions of the developing rat brain. Pediat. Res. 3:407. Pediat. Res_ 31:898. Fish, 1., and M. Wmick. 1969. The effects of malnutrition Wmick, M., and P. Rosso. 1969. Head circumferences on regional growth of the developing rat brain_ Exp. and cellular growth of the brain in normal and maras- Neurol. 25:534. mic children. J. Pediat. 74:741.

100 Wmick, M., and P. Rosso. 1970. Malnutrition and cellu- and after weaning) gave by far the greatest effects on lar growth in the brain. In J. Masek, K. Osancova, and food behavior and food consumption. D. P. Cuthbertson [ed.] Proc. Eighth Int. Congr. Nutr., Prague, Sept. 1969. Excerpta Medica, Amster- Malnutrition and Exploratory Behavior of Rats In dam. this study, which was in three phases, attention was given Winick, M., and P. Rosso. In press. Malnutrition and to the type of restriction and the time during which the central nervous system development. In Proceedings restriction was imposed. In all three phases, a diet with a of NICHD Conference on Neuropsychological Meth- subnormal protein level was fed to rats in the preweaning ods for Assessment of Impaired Brain Function in the period. In the first phase, the preweaning period was fol- Malnourished Child, Palo Alto, Calif., June 1969. U.S. lowed by severe protein restriction. In the second, the Government Printing Office, Washington, D.C. period was followed by severe calorie restriction. In the Winick, M., P. Rosso, and J. Waterlow. 1970. Cellular third phase, there was no restriction after the preweaning growth of cerebrum, cerebellum and brain stem in period. normal and marasmic children. Exp. Neurol. 26:393. After 4 weeks, the animals with protein- and calorie- restricted diets (those in the first two phases of the ex- Effects of Malnutrition in Early Life periment) were rehabilitated with the basal diet. Protein Postnatal Nutrition and Adult Behavior toward Food restriction caused lower body weight and slower develop- A restriction in the quantity of food consumed by a rat ment of exploratory behavior, beginning with the 10th in the first 3 weeks of life determines not only growth day of life. After weaning, severe protein or calorie re- rate during that time but also subsequent growth charac- striction caused elevation of spontaneous activity. After teristics when unlimited food is available. Studies have nutritional rehabilitation, exploratory activity declined been made of food consumption and body composition rapidly in the treatment groups. After 10 to 12 weeks, in adult rats that have plateaued in body weight but dif- malnourished rats showed a low exploratory drive. Inter- fer in ultimate size because of nutritional restriction post- pretation of a study such as this requires separate analy- natally. The study includes a detailed examination of sis of different characteristics of behavior. behavior toward food. Beginning at parturition, female rats were fed a 12- Malnutrition in Avoidance Conditioning of Rats The percent protein diet during the nursing period (3 weeks). study design described under "Malnutrition and Explor- The pups at weaning were divided into two groups, one atory Behavior of Rats," above, was followed in this receiving a 5-percent protein diet that prevented weight study. Rats were subjected to avoidance conditioning at gain in 4 weeks and the other receiving a restricted pro- 12 weeks of age. They learned to jump on a vertical tein diet that prevented increase in body weight. After screen when a conditioned stimulus was presented for 4 weeks all animals were given a normal controlled diet 10 seconds, after which unconditioned stimuli (electrical ad libitum. At 6 months of age restricted rats were shocks) were presented. Latent periods and spontaneous smaller than controls and consumed larger amounts of activity were recorded. There were no significant differ- diet per unit of body weight. Results indicate that post- ences between control and previously malnourished rats natal nutritional stunting of adult body size is not neces- in the learning rate expressed by the duration of latent sarily correlated with behavioral development. Behavioral periods, but striking behavioral disturbances developed abnormalities related to nutritional deprivations, how- in groups of rats that had been restricted both before and ever, are accompanied by a stunting of body size. Differ- after weaning. Behavioral abnormalities were manifested ences in eating habits were observed. The postnatally as inadequate stereotyped movements and inability to de- protein-restricted male animals exhibited significantly lay or to extinguish the fixed conditioned reactions. Ani- greater food spillage than the calorie-restricted or control mals restricted both before and after weaning showed groups. No difference in food consumption or food spill- signs of disturbance; those restricted only in the prewean- age was observed in female rats. The increased food spill- ing period did not show signs of disturbance. age that was observed in the early malnourished groups These studies demonstrate that the behavior of rats of animals was seen only when the animali were made can be affected not only by changes in the nutritional hungry. Hunger was induced by feeding them 1 hour per state during the preweaning period but also by changes day. If food was provided ad libitum, no differences in in the quantity or quality of the diet after weaning. food spillage were observed. Furthermore, this is not just A greater effect on behavior is exerted when restrictions a matter of calorie restriction but is due to calorie re- are imposed both before and after weaning. striction (restricted milk intake during suckling) followed by a low protein diet for a period of at least 4 weeks. Investigator: Richard H. Barnes, Graduate School of Nu- This combination of two forms of malnutrition (before trition, Cornell University, Ithaca, N.Y. 14850

101 Endemic Goiter in Latin America aged 1 to 5 months when they supplied 100 percent of The widespread occurrence of endemic goiter in Ecua- the total daily dietary protein. dor and Peru has led us to examine causes of the condi- Continuing studies on other sources of protein involve tion and practical therapeutic measures that may be ef- detoxification of rapeseed meal for both animal and hu- fective in improving iodine nutrition. The use of iodate man consumption. In view of the ready supply of rape- as a food supplement has been effective. We continue to seed, it is impoitant to provide a rapeseed meal protein look for the effect of supplementation on the appearance concentrate suitable for human use. of cretinism in the newborn, developmental milestones in the newborn, and the appearance of thyroid disease in Investigators: C. 0. Chichester, Department of Food Re- the young as well as in more mature age groups. We are source Chemistry, University of Rhode Island, Kings- also attempting to relate general intelligence and per- ton, R.I. 02881 formance to treatment in cases of goiter. Fernando Monckenberg, Department of Pediatrics and Iodized oil has been used as a prophylactic feature Biochemistry, School of Medicine, University of Chile, and has proved feasible without significant undesirable Santiago, Chile side effects and has been effective in preventing endemic goiter. Food Composition Table for Use in East Asia This project is sponsored by the Department of Health, Investigators: John B. Stanbury, Department of Nutrition Education, and Welfare and by the Food and Agriculture and Food Science, Massachusetts Institute of Tech- Organization. nology, Cambridge, Mass. 02139 The countries represented in the information on food Rodrigo Fierro, Ignacio Ramirez, and Eduardo Estrella, composition are Burma, Thailand, Laos, South Vietnam, Escuela Politecnica Nacional, Departmento de Radio- isotopos, Apartado 2759, Quito, Ecuador Cambodia, Malaysia, Singapore, Indonesia, Hong Kong, John T. Dunn, Department of Intemal Medicine, Uni- Philippines, Taiwan, Korea, and Japan. Data are being versity of Virginia School of Medicine, Charlottesville, collected on the proximate composition and on minerals, Va. 22901 vitamins, amino acids, fatty acids, and trace elements that have nutrient value. Information is accumulated Philip Dodge, Department of Pediatrics, Washington Uni- versity School of Medicine, 500 South Kingshighway, through communication with key investigators, with li- St. Louis, Mo. 63110 braries, and with other sources in the countries of inter- Eduardo Pretell, Instituto de Investigaciones de la Altura, est. Food data are being tabulated in a format previously Departmento de Endocrinologia, Apartado 6083, used for the African Food Composition Table, published Lima, Peru in both English and French. It is hoped that any labora- Andries Querido, Dukzigt Hospital, Dr. Molewaterplein tory having accurate information on the composition of 41, Rotterdam, Holland foods of East Asian countries will communicate such information to the principal investigators. Evaluation of Fish Protein Concentrate Investigators: W. T. Wu Leung, Nutrition Program, Health In Chile, as in most developing countries, one of the Services and Mental Health Administration, US. De- most serious sociomedical problems is chronic infant mar- partrrient of Heeth, Education, and Welfare, 9650 nutrition, which retards development, growth, and psy- Rockville Pike, Bethesda, Md. 20014 chological performance. Efforts to improve protein in- Mrs. R. R. Butrum, American Institute of Nutrition, take have been directed to utilization of fish protein 9650 Rockville Pike, Bethesda, Md. 20014 concentrate in preschool children's food. This work has been done in a single site near Santiago. It will be extended to other areas. Hair Root Morphology and Protein Synthesis in Disease Chemical composition, biological quality of the protein, and toxicity were determined for fish protein con- Changes in hair follicles are sensitive indicators of pro- centrate, sunflower presscake meal, and foods supple- tein synthesis under conditions of protein inadequacy. mented with these concentrates. Acceptability and Three types of studies have been conducted. They were tolerance were tested in human trials. The fish protein concerned with (1) experimental protein deprivation ard concentrate improved the biological quality of cereal- repletion of young adults in a metabolic ward, (2) grossly based foods and promoted normal growth in infants. obese men who were being fasted for an 8-week period, Mixtures containing fish protein concentrate and sun- and (3) children recovering from both forms of protein- flower presscake meal showed a high net protein utiliza- calorie malnutrition in South America, and children with tion in the rat and produced normal growth in infants mild to moderate PCM in the Caribbean area.

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1 03 In two separate studies, volunteers fed protein-free Improved Methods of Extracting and diets demonstrated significant hair root changes at the Utilizing Plant Proteins same time that urinary nitrogen levels had plateaued. Fermented foods are important components of diets Serum albumin and total serum protein values were nor- in many parts of the world. A study of the value of fer- mal at this time. When protein was added to the diet, the mentation in improving the nutritive value of food has changes in hair roots reverted to normal. In children who been conducted. Fermentation does not generally im- have had either acute or chronic protein-calorie malnu- prove the nutritive value of the protein but mayincrease trition, hair root recovery parallels closely the DNA- vitamin content. Digestibility for humans of certain fer- protein values from muscle biopsy samples. In the Carib- mented foods (e.g., tempeh, ontjom, and bongkrek) is bean studies of Negro preschool children, reduction in improved and the keeping quality is generally increased. weight for age is accompanied by a reduction in hair root Cooking time is often decr.eased. Organoleptic character- diameter. A study was conducted in Guatemala com- istics are generally improved. The wholesomeness of fer- paring eight parameters that have been suggested for the mented foods is generally as good as or better than that early recognition of malnutrit:on. The results of this of the ingredients from which they are produced_ When study are being calculated. aflatoxin-containing ingredients are used, there may be a hazard to health. Investigators: Robert B. Bradfield, Clinical Professor of The effect of fermentation was studied to improve the Human Nutrition, Department of Nutritional Sciences, efficiency of the process for extracting oil and protein University of California, Berkeley, Calif. 94720 from coconut meat. Coconuts available commercially D. B. Jelliffe, Professor of Community Nutrition, Univer- vary widely in maturity and length of storage and in re- sity of the West Indies, Mona Post Office, Kingston, sponse to fermentation processing. Sixty percent ofindi- Jamaica vidual coconuts showed a breaking of emulsion with separation of oil and proteins when fermented under The following publications resulted from this work: controlled conditions. Conditions of grinding and range of dilution were critical for rapid fermentation. The or- Bradfield, R. B. 1971. Protein deprivation: comparative ganisrn Lactobacillus plantantm was most effective in response of hair roots, serum protein, and urinary breaking the emulsion to yield a high grade, clear oil and nitrogen. Amer. J. Clin. Nutr. 24:405-410. with precipitated protein that was easily recovered in Bradfield, R. B., and M. A. Bailey. 1969. Hair root re- relatively undenatured form. sponse to protein undernutrition. In Advances in biology of skin. IX. Hair growth. Proceedings of a sym- Investigators: Keith H. Steinkraus, L. Ross Hackler, and posium at University of Oregon MedicA School, 1967. A. G. van Veen, New York State Agricultural Experi- Pergamon Press, New York. ment Station, Cornell University, Geneva, N.Y. 14456 Bradfield, R. B., M. A. Bailey, and A. Cordano. 1968. Hair-root changes in Andean Indian children during Increased Protein Supplies through marasmic kwashiorkor. Preliminary communication. Peanut Improvement Lancet, Nov. 30, 1968, 13. 1169-1170. Bradfield, R. B., M. A. Bailey, and S. Margen. 1967. Mor- Cooperative research in the United States and in India phological changes in human scalp hair roots during indicates that the protein content of peanut seed could deprivation of protein. Science 157:438-439. be increased by breeding. The average protein content of Bradfield, R. B., A. Cordano, and G. G. Graham. 1969. sound mature seed of a large number of diverse peanut Hair-root adaptation to marasmus in Andean Indian genotypes from the two countries ranged from 23.6 to children. Lancet, Dec. 27, 1969, p. 1395-1397. 30.5 percent when the genotypes were grown at Ludhi- Bradfield, R. B., and E. F. P. Jelliffe. 1970. Early assess- ana, India, or at Tifton, Georgia. The possibilityof in- ment of malnutrition. Nature 225:283-284. creasing the protein content of seed by breeding appears Bradfield, R. B., E. F. P. Jelliffe, and J_ Neill, Jr. 1970. more favorable for peanuts of the Virginia botanical type A comparison of hair root morphology and arm cir- than for the Spanish type. cumference as field tests of protein-calorie mahiutri- An obvious way to increase protein supplies is to use tion. J. Trop. Pediat. 16:195. higher-yielding varieties. When 100 diverse genotypes Bradfield, R. B., T. Yee, and J. M. Baertl. 1969. Hair zinc from India and 100 diverse genotypes from the U.S. col- levels of Andean Indian children during protein-calorie lection were grown at Ludhiana, India, from 1964 to malnutrition. Amer. J. Clin. Nutr. 22:1349-1353. 1966 in small-scale replicated tests, several Indian and U.S. genotypes substantially outyielded standard checks_ The most promiging of these have been included in larger- scale rests at Ludhiana. In general, the U.S. genotypes

103 104 that gave superior yields at Ludhiana show a high yield Hammons, R. 0. 1970. Spancrossa new peanut variety. potential when grown in adapted areas in the United Georgia Agr. Exp. Sta. Rep. 76. States. Hammons, R. 0. 1970. The Tifspan peanut variety. In cooperative research with the Georgia and Okla- Georgia Agr. Exp. Sta. Rep. 77. homa Agricultural Experiment Stations, three new productive Spanish peanuts of medium-sized seed were re- Influence of Specific Nutritional Deficiency leased to growers in 1969 and 1970. These are Spanhoma, on the Genetic Expression of Anemia a pure-line selection from Argentina; Tifspan, selected from progeny of a cross between Argentine and Spanish In 1968 the investigators studied the general problem 18-38; and Spancross, selected from progeny of a cross of multihemoglobins within erythrocytes. Observations between Argelitine and the wild annual, decumbent spe- made during that study suggested the feasibility of study- cies Arachis monticola. Argentine is a pure-line selection ing the effect of iron and folate deficiency on relative from Plant Introduction 121070, which was introduced synthetic rates of sickle and normal hemoglobins in pa- into the United States from Argentina in 1938. tients with sickle-cell trait. Little useful genetic resistance to pests has been found Later (1968-1969) the investigators and G. Milner studied the iron effect. Patients with sickle-cell trait were in cultivated peanuts. Certain related wild Arachis species are immune or highly resistant to several major pests, in- surveyed for relative proportions of Hb A and Hb S in cluding Cercospora leafspots, peanut rosette, peanut their erythrocytes. Bone marrow samples for culture were taken from iron-deficient subjects showing dispro- stunt virus, and peanut rust. Wild Arachis species with such characteristics cannot be crossed successfully with portional amounts of the two hemoglobins. The dispro- cultivated peanuts. Cooperative research is in progress portion was always in the direction of low Hb S. Iron with the North Carolina and Oklahoma Agricultural Ex- was added to these marrow cultures to determine periment Stations to correct this and make possible the whether it would correct the disproportional synthesis. incorporation of resistance or immunity to pests into im- Since no consistent effect was noted, it is probable that proved commercial varieties. Such a development could iron deficiency affects synthesis at a less differentiated result in enhanced production where there are pests for site than that tested in these studies. This work suggests which no economic control is now available and could certain aspects of biochemical genetics for future work. help sustain production levels if certain pesticides now used to control pests should become unavailable as a re- investigators: John F. Bertles, St. Luke's Hospital Center, sult of restrictions on their use or for other reasons. Amsterdam Avenue at 114th Street, New York, N.Y. Peanut genotypes with pods and seeds that resist in- 10025 vasion by Aspergillus flavus or A. parasiticus are being Paul F. A. Milner, Pathology Department, University of sought in cooperative research with the Alabama, Florida, the West Indies, Kingston, Jamaica and Georgia Agricultural Experiment Stations. Prelimi- nary results indicate that seeds of certain genotypes are Malnutrition in the Middle East more resistant to invasion by A. flavus than others. If Studies on the requirement of human infants for folic productive varieties with a high level of resistance to acid have been completed on 10 subjects maintained for aflatoxin-producing molds can be developed, millions of extended periods on a diet deficient in folic acid. Initial tons of high-protein peanut cake or meal now unsuitable obser;ations indicate that between 20 and 40 pg of folic for food because of contamination by aflatoxins could acid daily approaches the amount necessary to meet the be available for human consumption. requirements of an infant for hemopoiesis. There is also Exchange of peanut genotypes and production know- evidence that frequently occurring deficiencies of folic how is continuing with India and other countries. In the acid, protein, and iron are causes of the anemia of United States, research is continuing on the effect of kwashiorkor in Cairo. genotype and production environment on the protein A survey of the selenium and chromium content of content of mature peanut seed. In our peanut variety im- 85 foods indigenous to the Middle East has been com- provement research, special emphasis is placed on in- pleted. The foods include pulses and seed, cereals and creased yield potential, resistance to pests, and enhanced grains, vegetables, fruits, dairy products, meat, and eggs. processing and nutritional quality. In the Cairo area, selenium and chromium levels have been determined in plasma and erythrocytes of healthy Coordinator: W. K. Bailey, Plant Science Research Di- children and of children with kwashiorkor. Values for vision, Agricultural Research Service, Beltsville, Md. selenium in the blood of adults have been obtained by 20705 activation analyses. Some of these values are lower than values reported in the United States. The selenium con- The following publications resulted from this work: tent of serum and erythrocytes of healthy children in

104 na Cairo is significantly greater than that of children with to Group I at 6 weeks elicited a second reticulocyte re- protein-calorie malnutrition. sponse. Investigations into ways of improving the ability of Children with protein-calorie malnutrition in northern the Egyptian sandrat (Psammoruys obesus) to survive in Thailand present the classic syndromes of kwashiorkor captivity have been successful, and metabolic studies are and marasmus. The limiting nutrients for hemopoiesis ap- under way on this species. pear to be protein and iron. The delayed reticulocyte re- Evidence pertaining to the epidemiology of xeroph- sponse observed necessitates careful interpretation of the thalrnia and hypovitaminosis A in the Middle East has effect of any agentgivenas treatment. The results sug- been reported_ gest that alpha-tocopherol is not a primary nutrient for An interpretative monographic review on nutrition in hemopoiesis. the Middle East is being prepared for publication, and a second volume on the history and cultural role of food Investigators: Robert E. Olson, Donald Allen, Max N. in the Middle East is near completion. Horwitt, Franz Simmersbach, Jo Ann Whitaker, Charles H. Tan, and David Morehead, St. Louis Uni- Investigator: William J. Darby, Division of Nutrition, versity School of Medicine, 1402 South Grand Boule- Vanderbilt University, Nashville, Tenn. 37203 vard, St. Louis, Mo. 63104

Multiple Etiology of Nutritional Anemias Mycotoxin Contamination of Food and of Thailand Foodstuffs in Southeast Asia The first study of anemia and malnutrition in Chiang Aflatoxins have been .dentified chemically in varying Mai, Thailand, has been completed. Its objective was to amounts and frequencie_ 'n market samples of many determine whether vitamin E has a unique role in the kinds of Thai foods. The commodity contaminated at hemopoiesis of children with protein-calorie malnutrition. highest levels and frequencies is peanuts. Other principal Seventy-five children with protein-calorie malnutrition, vectors of aflatoxins are corn and various legumes. Rice, 6 months to 4 years of age, were studied. The children the principal dietary staple, is rarely contaminated. Afia- displayed anemia with hemoglobin levels lower than 9.8 g toxin ingestion is being measured directly by dietary sur- per 100 ml. They were underweight and of less than ex- vey techniques in which plate samples of foods are ana- pected height for age. lyzed and ingestion is calculated. Although aflatoxins are The most prominent vitamin deficiency was that of found in some instances, the level and frequency of ex- vitamin A. Thiamin and riboflavin deficiencies were the posure are much lower than market sample data had most obvious of the B-complex. Mortality rate declined indicated. from 25 percent the first 6 months to 4 percent the last Numerous samples of foods became highly toxic to 6 months principally as a result of a diagnostic and thera- rats when the endogenous molds were permitted to grow peutic attack on infections. under laboratory conditions. About 50 isolates of various The children were divided into five groups. Groups I genera of molds were found to be toxin-producing when and IB were fed the house diet for treatment of protein- grown in natural substrates. Systematic attempts are calorie malnutrition. This diet was adequate in protein being made to isolate and identify the toxic substances. and calories and in vitamins and minerals except for iron, It is not known whether these mycotoxins occur in hu- tocopherol, and possibly folate. Groups II, III, and 1116 man foods or whether they are important as hazards to were fed a diet low in tocopherol but adequate in all public health. other nutrients. Vitamin E was administered to Group I The ingestion of aflatoxin and the incidence of pri- after 6 weeks, Group III after 3 weeks, and Group 1116 mary liver cancer are being me.:sured in two parts of after 6 weeks. Thailand in which the incidences of apparent liver can- Children in all groups grew comparably over the 6- cer are very different. It is anticipated that these data week period, and serum proteins regenerated at the same will eventually permit a statistical evaluation of the pos- rate in all groups. sible relation between exposure to aflatoxin and inci- Those in Groups I and IB had poor initial hemopoietic dence of liver cancer. response and no increase in hemoglobin, owing to iron Aflatoxins (and possibly other mycotoxins) may be limitation. Administration of vitamin E after 6 weeks involved in the etiology of an acute toxicity syndrome had no effect. Children in Groups II, III, and 1116 had a in children. good initial hemopoietic response, with a reticulocytosis of 9 percent lasting 5 weeks and increased hemoglobin of Investigators: G. N. Wogan and R. C. Shank, Department 1.5 g per 100 ml. Administration of tocopherol to Groups of Nutrition and Food Science, Massachusetts Insti- III and "6 elicited no response. Administration of iron tute of Technology, Cambridge, Mass. 02139

105 C. M. Christensen, Department of Plant Pathology, Uni- villages in northeastern Thailand. Emphasis was placed versity of Minnesota, Minneapolis, Minn. 55812 on nutritional status of children, dietary habits and pat- Natth Bhamarapravati, University of Medical Sciences, terns, and other aspects of living conditions. Information Bangkok, Thailand was collected by distributing questionnaires and by making visits during which preschool children were given Nutrition and Infections in India complete medical examinations. The information re- In a study of nutrition and infections in India, illness vealed general poverty, low educational level on the part and growth in children under 3 years of age living in of the children, and varying degrees of malnutrition in Punjabi villages are being measured. The effects of com- each of the villages. plete medical care plus nutrition care, nutrition care Although the financial support for this work, fur- alone, and medical care alone are being compared. In nu- nished by the National Institutes of Health, was termi- trition care, mothers are advised about weaning prac- nated in 1969, Dr. Dradjat D. Prawiranegara in Indonesia tices, and the use of supplements in feeding stations is and Dr. Aree Valyasevi in Thailand have been able to confined to children who show a failure to grow by continue the study of communities with the assistance monthly weight measurements. Preliminary results show of volunteei workers. The workers, who live in the com- a synergistic effect of the combined complete medical munities, are trained in the fundamentals of good eco- care plus nutrition care as compared with medical care logical practice. Their contribution has led the Indone- alone. Continued observation on the influence of nutri- sian government to recognize the importance of tion care alone is necessary since our results so far are volunteers. The government's new 5-year plan includes inconclusive. the use of some 70 volunteers in its community improvement effort. Investigators: Carl E. Taylor and William A. Reinke, Volunteers were engaged in a similar undertaking in School of Hygiene and Public Health, The Johns East Java, Indonesia. The volunteers, most of whom Hopkins University, 615 North Wolfe Street, Balti- were college graduates who had studied human ecology, more, Md. 21205 were 'sent into a community in the city of Bozor. Each Arnfried Kielmann (Field Director) and I. S. Uberoi, lived with a family and through this relationship was able Narangwal Village Ludhiana District, Punjab, India to demonstrate to the community the value of improved living practices. P. N. Wahi and V. Ramalingaswami, Indian Council of Medical Research, Ansari Nagar, Post Box 494, New Delhi, India Principal investigators: Paul GS-forgy, University of Pennsylvania, Philadelphia General Hospital, Phila- Nutrition Research in Thailand and Indonesia delphia, Pa. 19104 Aree Valyasevi, Dean and Professor of Pediatrics, In the northern and northeastern provinces of Ramathibodi Hospital, Bangkok, Thailand Thailand, a serious medical problem, especially in rural Robert Van Reen, U.S. Naval Medical Research Institute, communities, is the frequent occurrence of bladder- Bethesda, Md. 20014 stone disease in children. Biochemical examinations were Dradjat D. Prawiranegara, Professor of Nutrition, Uni- given to village children and to urban children, and the versity of Indonesia, and Assistant Dean, School of results were compared. It was found that village children Public Health, Jakarta, Indonesia excreted significantly lower levels of inorganic phosphate and inorganic sulfate and that the incidence of oxal- Nutritional Anemia in East Asia crystalluria was much lower. Administration of methionine to village children resulted in increased urinary iii- Iron deficiency anemia is widespread throughout the organic sulfate, whereas vitamin B6 supplementation had world. In addition, megaloblastic anemia is a problem in no effect. These results indicate an inadequate intake of world health_ The latter may result from any defect in protein. In the bladder-stone cases (rural children), sup- nucleoprotein synthesis, but in more than 95 percent of plementation with inorganic orthophosphate resulted in the cases it results from deficiency of folic acid or vita- an increase in urinary orthophosphate and pyrophos- min B12. Folate deficiency appears to occur among phate, and there was an associated reduction in urinary many of the economically disadvantaged people of the calcium and oxalate. Oxalcrystalluria was seen to dis- world, especially pregnant women_ A deficiency of vita- appear within 24 hours after supplementation. The con- min B12 is common among vegetarians, patients with clusion was that in the Thai provinces bladder-stone dis- sprue, and the elderly. These studies indicate that folate ease results chiefly from diets that are low both in and vitamin B12 deficiencies resulting in serious megalo- protein and in phosphate. blastic anemia occur widely and are of particular interest Human ecological studies have been conducted in 10 in East Asia.

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107; The World Health Organization has initiated collabora- Fernandez, N. A., J. C. Burgos, C. F. Asenjo, and I. R. tive studies of these diseases. Reference centers for the Rosa. 1969. Nutrition survey of two rural Puerto studies have been established at St. Bartholomew's Hos- Rican areas before and after a community improve- pital in London (under Professor David Mollin), at the ment program. Amer. J. Clin. Nutr. 22:1639-1651. Instituto Venezolano de Investigaciones Cientificas in Caracas (under Professor Miguel Layrisse), and at the Relationship of Diet to the Performance of the Kuala University Hospital of the University of Malaya in Combat Soldier Lumpur (under Professor Kam-Seng Lau and Dr. Herbert). This program is designed to help each countrydelineate The marked incidence and severity of acute mountain the incidence, etiology,.and treatment of nutritional sickness (AMS) resulting from abrupt altitude exposure of anemia. is a major problem in maintaining maximal efficiency Studies are continuing to examine the possibility of troops, and could seriously impair mllitaryperformance an etiologic infectious agent intropical sprue. This dis- and operations. In past studies the severity of AMS ease, in which anemia is a common finding,affects nearly symptoms were gradually increased, reaching theirmaxi- 100 million people in East Asia. mum at 36 hours of acute altitude exposure at4,300 meters. In two later studies, the AMS were of shortdura- Investigator: Victor Herbert, The Mount Sinai School of tion, reaching their maximum at 6-10 hours. In bothof Medicine, 100th Street at Fifth Avenue, New York, these later studies, the men were physically conditioned N.Y. 10029 prior to altitude exposure and were consuming normal and high carbohydrate diets. The following publication resulted from this work: Anorexia has been reported in previous studies by our laboratory and has usually been accompanied by nega- Herbert, V. 1968. Megaloblastic anemia as a pioblem in tive nitrogen and water balances. In one study, it was ob- world health. Amer. J. Clin. Nutr. 21:1115-1120. served that, with some motivation, the daily food intake can be maintained after abrupt altitude exposure.Under these conditions, one observes positive nitrogen and min- Nutritional Studies in Puerto Rico eral balances, greatly reduced body weight losses, normal A survey in the slum area of Juana Matos, Catatio, blood electrolytes, and normal fasting glucose levelsand Puerto Rico, was conducted as part of a continuing pro- glucose tolerance curves. It appears that many of the gram in Puerto Rico. The area is across thebay from biochemical changes previously attributed to hypoxia at San Juan. It had a population of 2,655, divided into 5114 altitude may be partly due to anorexia and the subse- families. The survey included clinical examinations quent caloric deficit. (1,796 persons), biochemical tests (910 persons), and Body hypohydration has been shown to occur at alti- collection of dietary data (59 families). tude during acute exposure. Although extracellular fluid Annual income in half the families was $1,000 to is practically unchanged, total body water andintracellu- $3,000 and the general educational level was low. lar waters are both significantly decreased. Otherbody The diet was generally below the recommended al- fluid changes include a negative water balance, signifi- lowances for calories, calcium, vitamin A, vitamin C, and cantly decreased blood and plasma volumes, and a nat- riboflavin. The prevalence of observable nutritional de- ural diuresis. ficiency was low. Retardation of growth, as expressed in Heavy physical activity and physical conditioning height and weight for age, and considerable obesity prior to acute high-altitude exposure can greatly reduce among women were observed. the severity of acute mountain sickness (AMS).During Hematologic findings showed a low incidence of acute altitude exposure, there appears to be aminimal hypochromic anemias or low plasma protein levels. carbohydrate requirement. One usually observes anorexia Plasma ascorbic acid was adequate in all subjects. Uri- (25 to 40 percent decrease in food intake) during acute nary excretions of riboflavin and niacin indicated asub- altitude exposure, and under these conditions the daily stantial degree of deficiency. Intestinal infestation with carbohydrate intake should be at least 320 g. It appears helminth parasites was prevalent. that body hypohydration and a naturally occurring cold diuresis are adaptive mechanisms for high-altitude Investigators: Nelson A. Fernandez, Jose C. Burgos, and exposure. Conrado F. Asenjo, Department of Biochemistry and Nutrition, School of Medicine, University of Puerto Investigators: C. Frank Consolazio, H. L. Johnson, H. J. Rico, San Juan, Puerto Rico 00905 Krzywicki, and T. A. Daws, U.S. Army Medical Re- search and Nutrition Laboratory, Fitzsimons General The following publication resulted from this work: Hospital, Denver, Colo. 80240

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108' The following publication resulted from this work: TEMPERATE ECOLOGY

Consolazio, C. F., L. 0. Matoush, H. L. Johnson,H.J. Good-quality protein foods are abundant in the Krzywicki, T. A. Daws, and G. J. Isaac. 1969. Effect United States, Canada, Western Europe, and other coun- of high carbohydrate diets in performance and clinical tries in the Temperate Zone. Nevertheless, rescarch is symptomatology after rapid ascent to high altitude. needed to improve the food produced for our own peo- Fed. Proc. 28:937-943. ple and for export. Genetic research to improve the protein quality and yleld of cereal foods, and research on all Role of Zinc Deficiehcy in Growth Retardation other types of potential natural sources of protein, must continue. This study was designed to determine whether zinc Most of the nutrition research dealing with problems deficiency was the cause of pronounced growth retarda- in the Temperate Zone is relevant to tropical countries tion and delayed or absent sexual maturation in subjects also. However, the investigations described here relate from villages near Shinaz, Iran. Seventeen "dwarfs," all primarily to local problems and, in general, are con- aged 19 or 20, who had been rejected by an Iranian ducted in the United States. Army induction center ageed to stay in the hospital for a year. They were divided into three groups. Group I received a well-balanced hospital diet containing ample ani- Diet and Fat Metabolism mal protein. Group II received the same diet plus a cap- Although the carbohydrates have long been con- sule of zinc sulfate daily. Group III received the hospital sidered of nutritional importance, chiefly as a source of diet for 6 months and the diet plus zinc sulfate for the calories, there is increasing interest in the individual car- next 6 months. Analysis of rate of growth and onset of bohydrates because of possible differences in their ef- sexual functioning revealed that the diet alone resulted fect on lipid metabolism. Investigations on the influence in gradual growth and sexual maturation but that the ad- of several types of dietary carbohydrate on lifespan and dition of zinc sulfate resulted in greatly enhanced growth lipid metabolism have been conducted with two strains and development (P < .001). of experimental rats. Cholesterol and noncholesterol It is concluded that zinc deficiency occurs in man and lipids were generally higher in livers of both strains when produces abnormalities similar to those seen in experi- diets containing 2 5 percent of cooked egg and sucrose mental zinc deficiency of animals. were fed than when the carbohydrate was dextrose or The cause of this zinc depletion is thought to be the cornstarch. Carbohydrate differences were much more diet consumed by villagers, which consists largely of un- marked in a strain of rats developed in this laboratory leavened bread. This has a high content of phytate, (BHE) than in a strain of Wistar rats. Renal disease, the which binds zinc, making it unavailable. The degree of chief cause of death in the BHE rat, was accelerated by this problem is being investigated in Iran. Various breads the diet containing sucrose. Degenerative changes in the are being analyzed for phytate and the effect of fermen- kidney were rarely seen in the Wistar rat, and survival tation on destroying phytate_ Studies of zinc and nitro- was not significantly influenced by type of diet. Studies gen balance are being conducted in the metabolic ward with three other commonly used strains of rats, directed of Nemazee Hospital. Seven of the first eight apparently toward establishing the extent to which fructose may be healthy village adults were in strongly positive zinc bal- contributing to the metabolic response observed with su- ance, which indicates a deficient state. There was a tend- crose, have been completed but have not yet been ency toward negative nitrogen balance, which suggests evaluated. impairment of protein synthesis in the zinc-deficient state. Investigators: Mildred Adams and Duane Benton, Human Investigators: James A. Halstead, Consultant in Clinical Nutrition Research Division, Agricultural Research Nutrition, Trace Element Research Laboratory, Vet- Service, Beltsville, Md. 20705 erans Administration Hospital, Washington, D.C. 20422 The following publications resulted from this work: John G. Reinhold, Director, Pahlavi Nutrition Research Project, Pahlavi University, Shinaz, Iran Durand, A. M. A., M. Fisher, and M. Adams. 1968. The James C. Smith, Jr., Chief, Trace Element Research influence of type of dietary carbohydrate: effecton Laboratory, Veterans Administration Hospital, histological fmdings in two strains of rats. Arch. Washington, D.C. 20422 Pathol. 85:318-324. Lakshmanan, F. L., E. M. Schuster, and M. Adams. 1967. Effect of dietary carbohydrate on the serum protein

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109 components of two strains of rats. J. Nutr. 93: Hodges, R. E., E. M. Baker, J. Hood, I. E. Sauberlich, 117-125. and S. C. March. 1969. Experimental scurvy in man. Taylor, D. D., E. S. Conway, E. M. Schuster, and M. Amer. J. Clin. Nutr. 22:535-548. Adams. 1967. Influence of dietary carbohydrates on Hood, J., and R. E. Hodges. 1969. Ocular lesions in liver content and on serum lipids in relation to age scurvy. Amer. J. Clin. Nutr. 22:559-567. and strain of rat. J. Nutr. 9:275-282. Human Iodine Requirements and Substances in Experimental Scurvy in Man Foods That Affect Thyroid Function There is considerable difference of opinion regarding Although goiter has been found to occur generally in the amount of vitamin C necessary in-the diet of normal areas where iodine intake is low, in some areas amajority adults. Some authorities believe that tissue saturation is of the population show no clinical evidence of abnormal beneficial whereas others believe that saturation is not thyroid function in spite of subnormal levels and low necessary in the interest of health. The objectives of this iodine intakes. To study the possible mechanism in- project were to measure, by means of carbon 14 labeled volved, the distribution of iodoamino acids and iodopro- I-ascorbic acid, the body pool of vitamin C and to study teins in the thyroid glands of goats was investigated. its rate of depletion during ascorbic acid deprivation. Thyroids were obtained from goats from an endemic Six healthy men were fed a diet devoid of vitamin C goiter area and from an area free from endemic goiter. but adecuate in all other nutrients. Mild scurvy devel- Previous studies had shown that the thyroids of goats oped in the third month of deprivation. Clinical symp- from an endemic goiter area had pathophysiological toms of scurvy were present when the pool size of the changes similar to those found in human thyroids from vitamin had decreased to about 300 mg and the rate of the same area. Thyroid glands of goats living in an area catabolism of vitamin C was less than 9 mg daily. of severe environmental deficiency of iodine showed Urinary excretion of ascorbic acid ceased by the 23rd higher mono-iodotyrosine/di-iodotyrosine and tri- day of deprivation and was again detected after the body iodothyronine/thyroxine ratios than the glands of those pool of ascorbic acid was partly repleted. A daily intake living in an area of iodine abundance. There were also of 6.5 mg was sufficient to cause slow disappearance of differences in the incorporation of 1311 in the iodopro- clinical signs of scurvy in one subject. teins of the thyroids. Despite the recorded rarity of ocular vascular lesions The OZT (5-vinyl-I, 2-oxazolidine-thione), ITC (ise- in human scurvy, conjunctival lesions appeared in five of thiocyanate), and TC (thioCyanate) content of seven nine subjects with induced scurvy. The lesions ranged vegetables has been determined and the effect of these from minute bulbar conjunctival hemorrhage to a large substances on the thyroids of chickens and rabbits has subconjunetival hemorrhage accompanied by palpebral been studied. The OZT was found to increase the 1131 petechial hemorrhages and intense congestion of the con- uptake by thc thyroid. This was proportional to the in- junctival vessels. crease in size and was not overcome byiodide. The ITC The data reported and data obtained in a subsequent and TC compounds reduced the 1131 uptake. Rapeseed study, the results of which have not yet been published, meal had a high level of OZT and significant levels of are in agreement with the results of the MedicalResearch ITC and TC, and it was active in thyroid enlargement. Council Sheffield study, which indicated that a daily Cows fed rapeseed meal did rhave OZT in the milk, dose of 10 mg of ascorbic acid is sufficient to cure scur- but significant levels of TC were found. Milk from cows vy in man.* The results also suggest thzt 30 mg is an ade- fed rapeseed meal did not produce a good quality of quate daily allowance of ascorbic acid. cheese_ Some substances in the milk apparently inter- Investigator: Robert E. Hodges, Department of Internal fered in the ripening of the cheese. The levels of goitro- Medicine, University Hospitals, University of Iowa, gens found in vegetables are not considered great enough Iowa City, Iowa 52240 to produce goiter in humans consuming normal amounts. The following publications resulted from this work: Investigators: Ruth Leverton, Mildred Adams, and Duane A. Benton, Human Nutrition Research Divi- Baker, E. M., R. E. Hodges, and J. Hood. 1969. Metab- sion, Agricultural Research Service, Beltsville, Md. olism of ascorbic-1-14C acid in experimental human 20705 scurvy. Amer. J. Clin. Nutr. 22:549-558. Nutritional Status of Preschool Children in the *Medical Research Council. 1953. Vitamin C requirement of hu- United States man adults: a report by the Vitamin C Subcommittee of the Ac- cessory Food Factors Committee. No. 280. Compiled by W. In view of reports of serious malnutrition among Bartley, H. A. Krebs, and J. R. P. O'Brien. Medical Research Council, London. 179 p. children in southern states, particularly Mississippi, we

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1110 undertook a survey of 585 preschool children in coopera- ma vitamin A assay by cluorometry and use of a silicic tion with the Assissippi State Board of Health and the acid column technique. Clin. Chem. 16:766. University of Mississippi Medical Center. This was a pilot Owen, G. M., P. J. Garry, K. M. Kram. C. E. Nelsen, and study in preparation for a larger national study. Data in- J. M. Montalvo. 1969. Nutritional status of Mississippi dicated some restriction of caloric intake of children in preschool children. Amer. J. Clin. Nutr. 22:1444- low-income families, and the calcium and ascorbic acid 1458. intakes appeared inadequate. Also, it was found that Owen, G. M., P. J. Garry, A. H. Lubin, and K. M. most of the children sampled were ingeaing insufficient Kram. In press. Nutritional status of preschool chil- iron to meet their needs. Mild iron deficiency anemia dren: plasma vitamin A. J. Pediat. appears to be common, particularly among poor Owen, G. M., C. E. Nelsen, and P. J. Garry. In press. Nu- children. tritional status of preschool children: hemoglobin. In 1968 children in two countries in southern Ohio hematocrit and plasma iron values. J. Pediat. were examined for nutritional adequacy. It was con- Owen, G. M., C. E. Nelsen, K. M. Kram, and P. J. Garry. cluded that serious hunger of malnutrition is not wide- 1969. Nutritional status of preschool children in two spread in those counties. A number of the children were counties in southern Ohio. Ohio State Med. J. 65: smaller than average at the time of the study. Anemia oc- 809-814. curred at 2.5 times the rate at which it occurs on a national scale. About one fifth of the preschool children Nutritional Value of Wheat and Wheat Products had low levels of plasma ascorbic acid. Calcium intakes were low in a significant number. A more extensive eval- Wheat makes up more than 80 percent of the cereal uation of the hematological status in children indicates products consumed as food in the United States. A most that iron deficiency is not an uncommon occurrence comprehensive study in terms of numbers of nutrients among preschool children in the United States. The inci- determined and numbers of wheat products studied has dence is probably greatest among poor children. It ap- been conducted to provide up-to-date information on pears that a preschool child with anemia frequently is de- the nutrient contributions of wheat and wheat products. ficient in iron. Five hard wheats, four soft wheats, two durum wheats, The field studies in this project have been completed and the flours milled from these wheats have been ana- but analysis of the data is yet to be published. lyzed for four B vitamins, tocopherols, 15 fatty acids, 14 During the course of the population studies it was minerals, protein, and amino acids. The various forms of possible to refine the method for quantitating vitamin A the vitamins were measured, and changes that occurred in serum or plasma through the use of silicic acid column during milling and processing were determined. chromatography and fluorometry. A fluorescing inter- Only 5 of the 15 fatty acids were present in significant fering substance was identified, and a procedure for dif- amounts in the samples studied. Linolenic acid occurred ferentiating it from vitamin A was developed. Also inci- in the largest amounts. Individual samples from all re- dent to the large study was an attempt to identify pheno- gions of the country had essentially the same fatty acid types of serum cholinesterase. An automated method composition. About 10 percent of the tocopherols pres- has been described that measures serum cholinesterase ent in hard and soft wheats and 40 percent of those in activity and identifies the usual and atypical forms of durum wheat survived the milling and bleaching process. enzyme. Of the total tocopherol content, two thirds was in the form of epsilon tocopherol and less than 10 percent was Investigator: George M. Owen, Department of Pediatrics, in the alpha form. Children's Hospital, The Ohio State University, Major losses occurred in the B vitamins, thiamin, ribo- Columbus, Ohio 43205 flavin, niacin, and vitamin 136. Less than 20 percent re- mained after 60 percent extraction. The following publications resulted from this work: Changes in mineral content were significant Less than 25 percent of the iron, phosphorus, potassium, copper, Garry, P. J. 1971. A manual of an automated procedure magnesium, manganese, nickel, and zinc were retained. for measuring serum cholinesterase activity and identi- As expected, the lower the percentage of the extraction, fying enzyme variance. Differentiation by means of the lower the mineral content. This is important informa- tris and phosphate buffers. Clin. Chem. 17:192. tion in view of the essentiality of certain trace elements Garry, P. J. 1971. Serum cholinesterase variants: exam- and the variable avan.ability in foods. ination of several differential inhibitors, salts, and Consumption of diets containing 3% ounces of unen- buffers used to measure enzyme activity. Clin. Chem. riched flour daily would provide the following fraction 17: 181 of the Recommended Daily Allowance: 10 percent for Gany, P. J., J. D. Pollack, and G. M. Owen. 1970. Plas- thiamin, 2 percent for riboflavin, 8 percent for niacin,

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14' 4 percent for vitamin B6, 2 percent for calcium,less than The role of ascorbic acid in the mechanisms of anti- 10 percent for iron, 10 percent for magnesium, and 16 body synthesis has long been a matter of dispute.Ithas percent for phosphorus. been demonstrated that guinea pigs deficient in vitamin 86 produce less circulating diphtheria toxoidantibody. Investigators: W. A. Gortner and E. W. Toepfer, Human Severely scorbutic guinea pigs maintained on a highly Nutrition Research Division. Agricultural Research purified deficient diet were immunized with diphtheria Service, Beltsville, Md. 20705 toxoid. No deleterious effects on primary or secondary circulating antibody formation were observed in these The following publications resulted from this work: animals. Studies are being continued on the possible role of Eheart, J. F., and B. S. Mason. 1970. Nutrient composi- pantothenic acid in the functioning of individual tion of selected wheats and wheat products. V. Carbo- antibody-producing cells. This relates to a major interest hydrate. Cereal Chem. 47:715-719. in the role of pantothenic acid in protein synthesis and Inknen, J. A., and F. W. Quackenbush. 1969. Extractable transport. and "inbound" fatty acids in wheat and wheat products. Cereal Chem. 46:580-587. Investigator: A. E. Axelrod, Biochemistry Department, Polansky, M. M., and E. W. Toepfer. 1969. Nutrient School of Medicine, University of Pittsburgh, Pitts- composition of selected wheats and wheat products. burgh, Pa. 15213 IV. Vitamin B-6 components. Cereal Chem. 46: 664-674. The following publications resulted from this work: Slover, H. T., J. Lehmann, and R. J. Valis. 1969. Nu- trient composition of selected wheats and wheat Kumar, M., and A. E. Axelrod. 1968. Cellular antibody products. III. Tocopherols. Cereal Chem. 46:635-641. synthesis in vitamin B 6-deficient rats. J. Nutr. 96:53. Toepfer, E. W., E. M. Hewston, F. N. Hepburn, and J. H. Kumar, M., and A. E. Axelrod_ 1969. Circulating anti- Tulloss. 1969. Nutrient composition of selected body formation in scorbutic guinea pigs. J. Nutr. wheats and wheat products. I. Description of samples. 98:41. Cereal Chem. 46:560-567. Zook, E. G., F. E. Greene, and E. R. Morris. 1970. Nu- Protein Value of Plant Foods trient composition of selected wheats and wheat products. VI. Distribution of manganese, copper, The problem of meeting protein requirements with nickel, zinc, magnesium, lead, tin, cadmium, chro- plant proteins when high-quality animal proteins are not mium, and selenium as determined by atomic absorp- available or are not economically feasible is one of great tion spectroscopy and colorimetry. Cereal Chem. 47: concern. Current research is directed towardsupplying 720-731. information on selected aspects of this problem. Diets low in protein or containing proteins of poor quality were found to be associated with a decrease in Nutritive Factors in the Immunological Process brain glutamic dehydrogenase and decarboxylase and an The sivilficant role of vitamin B6 in the develop- impairment in psychological performance of the rat. In ment of immune processes in animals has been estab- contrast, mere caloric restriction prior to or after wean- lished by measuring change of serum antibodies in ani- ing had no such effect. The postweaning diet appeared mals deficient in vitamin B6. This is not unequivocal to be more important than the maternal diet fordevelop- proof since the serum antibody level reflects an equilib- ment of offspring, but the latter did affect breeding per- rium between the rate of synthes and release and the formance, birth weight, and weaning weight Dull and rate of destruction and excretion. There was a need to bright strains of rats have been developed by selective study the effect of vitamin deficiency on the individual breeding over four generations, and learning ability is discrete processes and on the metabolism of antigen be- being evaluated by selected measurements cf psycho- fore it reaches target tissue, such as the spleen. logical Cellular antibody synthesis was determined by the Investigations are under way to add to our informa- Jerne agar-plaque technique in normal rats and in rats de- tion on the possible supplementation of cereal diets deficient in vitamin B6. The decreased cellular immune re- ficient in lysine with legumes containing a high level of sponse observed in the deficient animals was independent this amino acid. The purpose is to find a way to formu- of the inanition and was restored to normal by adrnin- late low-cost meals for postweaning children. Prelimi- istering pyridoxine shortly before immunization. Ac- nary evaluations of various food combinations arebeing cumulation of antigen by the spleen was not impaired in made in growth studies of rats. A diet composed of ce- the deficient animals. real or millet with bengal gram and leafy vegetables

111 112 compared favorably with a 10 percent casein diet and Laboratory, University of Notre Dame, Notre Dame, was superior to a diet of bread enriched with vitamins, Ind. 46556 minerals, and lysine. Other combinations will be tested D. C. Anderson, Department of Psychology, Lobund and the most promising ones will be fed to young chil- Laboratory, University of Notre Dame, Notre Dame, dren to determine their nutritional response to such Ind. 46556 diets. Although there has been limited evidence suggesting Vitamin B6 Metabolism in Man that the protein efficiency ratio of tempeh, a fermented soybean product, is superior to that of the unfermented Not only is it important to develop more definitive soybean, recent research has failed to confirm this find- methods of assessing vitamin B6 nutriture in persons of ing. Protein efficiency ratios of freshly prepared or all ages, but more precise means of measuring the vita- stored tempeh and of unfermented soybeans were not min B6 requirement are needed. significantly different. Protein values of tempeh were im- A study recently completed demonstrates the forms proved when properly supplemented with lysine, methi- of vitamin 86 that are excreted by man with an intake onine, and threonine. The antioxidant isolated from of 1.66 mg daily. Under these conditions the B6 ex- tempeh was found to be ineffective as a substitute for creted is made up of 2/3 pyridoxal and 1/3 pyridox- vitamin E. amine. With a decrease of intake to 0.16 mg per day, the urinary vitamin B6 decreased rapidly. There appears to Investigators: Ruth M. Leverton, Mildred Adams, June be a specific effect of such depletion on the excretion of Kelsay, and E. W. Toepfer, Human Nutrition Research cystathionine and other methionine metabolites. Division, Agricultural Research Service, Beltsville, Md. Continued studies will investigate the vitamin B6 rela- 20705 tionships during pregnancy and during ingestion of oral contraceptives where abnormal tryptophan metabolism Relationship of Dietary Deficiency and is not an accurate indication of vitamin B6 deficiency in Impairment of Mental and Somatic women. Development Investigator: Hellen Linkswiler, Department of Nutri- With the aid of germfree and limited-flora animals, tional Sciences, College of Agxiculture and Life Sci- absolute levels of amino acids required for normal devel- ences, University of Wisconsin, Madison, Wis. 53706 opment, and the effect of intestinal microflora in changing those requirements, can be determined. Diet formula- tions, behavioral tests, and biochemical tests have been developed for determining interactions among early ARCTIC ECOLOGY amino acid deficier.,ies, intestinal microflora activities, and later behavioral changes in rats. Amino acid balances The subprogram on arctic ecology is sponsored jointly in defined diets have been improved, and solid-type by the United States and Canada and will be concerned amino acid diets have been found adequate for reproduc- with a series of Eskimo communities. It will consist of tion in germfree animals. A series of behavioral tests the following phases: measuring activity levels and learning capacity have been Clinical examinations for signs of specific defi- administered to young adult rats reared under germfree ciencies (serum iron, hemoglobin, serum proteins, blood or conventional conditions. Differences between the be- levels of vitamin C, folic acid, vitamin B12, and serum havioral responses of the two groups of rats were found lipids) in the initial studies, but the differences were reduced to Estimation of caloricintake by major food types nonsiglificant values in later tests in which closer con- and seasonal variations (dietary histories, diet analyses, trol -of other environmental variables was achievecL 24-hour urinary nitrogen, and excretion) Pancreatic and intestinal enzymes nave been assayed to Use and management of local food sources, distri- ensure that germfree animals fed defined diets have suf- bution of food in the community, and type of food ficiently normal levels to permit evaluation of the effects brought into the community of amino acid deficiencies and microflora activities on The subprogram is being prepared by H. E. Sauberlich, these critical enzymes. Liver enzymes involved in inter- U.S. Army Medical Research and Nutrition Laboratory, mediate carbohydrate metabolism have been studied to Fitzsirnons General Hospital, Denver, Colo. 80240, and assess the extent of general microflora effects. G. H. Beaton, University of Toronto, Toronto, Canada. Preliminary steps in the preparation are supported by Investigators: Bernard S. Wostmann, J. R. Pleasants, and the Medical Research and Development Command, U.S. B. S. Reddy, Department of Microbiology, Lobund Department of the Army.

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113 Nutrition Survey among Eskimosof and excretion data indicated adequate levels of water- soluble vitamins. Also, urinary nitrogen and calcium Wainwright, Alaska were quite high, indicating adequatedietary intakes of This nutrition survey wasconducted during January protein and calcium. The schoolchildren were provided a school lunch 1969 in cooperation with F. A.Milan, principal investi- each day, supplemented with vitamin tablets toprovide gator of the ecological study.Its purpose was to determine the nutritional status ofEskimos in the village of fat-soluble vitamins. The school has excellent facilities village of the for preparing and serving the lunches. Wainwright, which is the northernmost of United States, 90 miles from Barrowand 300 miles north A cooperative store provides significant amounts calories and nutrients for the village. Supplies are of the Arctic Circle. The Wainwdght population is 350(52 households), brought in by boat once a year, usually in September, The median age of males is 14.5 years,compared with and are adequate for most of the year. Items that provide ascorbic acid were found to be in good supply. 30.4 years for the male populationof the United States It appears that Wainwright Eskimos are able to main- as a whole. Twenty-seven percentare less than 6 yearsof tain themselves in a generally acceptable nutritional state age. Blood and urine samples wereobtained on one third despite their harsh environment and limited resources. of the population. Other appropriateobservations were made on the total population and itsfood resources, food Investigators: H. E. Sauberlich, W. Goad, and Y.F. practices, and health services. Blood samples were analyzed forhematocrit, hemo- Herman, U.S. Army Medical Research and Nutrition 1.-aboratory, Fitzsimons General Hospital, Denver. globin, total serum proteins, thewater-soluble vitamins, Colo. 80240 calcium, and nitrogen. U.S. Public Evaluation of the data obtained indicatesthat the F. A. Milan, Arctic Health Research Center, problem was anemia. This Health Service, College, Alaska 99701 most significant nutritional University of appears the result of aninadequate intake of dietary iron. P. Jamison, Department of Anthropology, The subjects studied had adequate serumprotein levels, Wisc on sin , Madison, Wis. 53706

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114 Chronobiology

The ubiquity and importance of biological rhythms ment are used. Measurements on healthy volunteers have become apparent during the last two decades. are taken while the subjects are under normal conditions Methods for detection, quantification, and analysis of for their culture. rhythms have been developed in the last decade. The rhythms of the heart, the brain, and respiration are taken for granted. Other more subtle rhythms of PHASE S-l!FTS circadian (about daily) frequency have been discovered. Body temperature and work fitness are examples of variables that follow a ercadian rhythm. Phase shifts of rhythms will be evaluated on healthy Circadian rhythms are often assumed to be caused volunteers_ Body rhythms will be measured first under by eating or work habits or other schedules. Studies normal conditions and then after intercontinental flights. have demonstrated that man, other animals, and plants A control group will take flights of the same duration continue to display circadian variations in the absence but will circle above the point of departure. These of known periodical changes in the environment. studies will be made on persons who have been subjected Franz Halberg, Department of Patholou, University to conditions routinely experienced by airpiane crews of Minnesota, is Program Director. and passengers, and the results will be compared with earlier fmdings, which suggest that delays of rhythms occur more rapidly than advances. Changes in mental and circulatory performance will be determined. Similar OBJECTIVES studies will be made on the effects of changes in work routine.

Objectives are: To develop a better definition of health by quan- OTHER STUDIES tifying normalcy according to rhythrnometiic criteria To measure the effects of shift work and trans- Projects concerned with body rhythms in cancer meridian flights on body rhythms chemotherapy, psychiatry, and other areas of clinical To diagnose maladaptation or early illness by medicine have been undertaken. rhythmometric analysis Additional studies are concerned with shift-work monitoring and arctic chronobiology.

RESEARCH PLANS RESEARCH PROGRESS

RHYTHMOMETRY STANDARDS RHYTHMOMETRY STANDARDS Comparable data will be obtained on selected human body rhythms. Variables selected are bodytemperature, Circadian rhythms of heart rate, oral temperature, pulse, blood pressure, certain performance tests, and and urinary hydroxycorticosteroids and potassium ex- various aspects of the urine (excretory rate, volume, cretion have been measured by investigators in various sodium and potassium content, and the concentration of parts of the world. The fact that all investigators found 17 hydroxycorticosteroids). Priority measurements in- similar rhythms indicates that the data can be usedas clude body temperature, performance, and urinary clinical standards. (See Halberg et al., 1969, in theap- sodium and potassium_ Standard methods ofmeasure- pended list of publications.)

114 115 PHASE SHIFTS ditional data on arctic chronobiology were collected for the IBP study of Eskimos (see p. 65). A study of the effects of transmeridian flights on man shows that the adjustmentsthat follow flights from east to west differ, in the same persons,from those that follow flights from west to east. (See Halberg, 1969, in APPENDIX: PUBLICATIONS the appended list of publications.) Bohlen, J. G., F. A. Milan, and F. Halberg. In press. Circumpolar chronobiology. Proc. IX Int. Cong. Anatomists, Leningrad, 1970. Halberg, F. 1969. Chronobiology. Annu. Rev. Physiol. 31:675- 725. OTHER STUDIES Halberg, F., W. Nelson, R. Doe, F. C. Bartter, and A. Reinberg. In press. Chronobiologie. J. Eur. Toxicol. Acrophase charts have been prepared for shift-work Halberg, F., J. Reinhardt, F. Bartter, C. De lea, R. Gordon, Halberg S. Wolff, A. Reinberg, J. Ghata, H. Hofmann, M. Halhuber, monitoring and for possible clinical use. (See R. Giinther, E. Knapp, J. C. Feta, and M. Garcia Sainz. 1969. et al., in press, in the appended list ofpublications.) Agreement in end points from circadian rhythmometry on Data on arctic chronobiology in relation to human healthy human beings living on different continents. performance on odd schedules have been collected. Ad- Experimenta 25:107-112.

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116 Atti.Fs' IF° IE Pa 1E, 10C_ IIE Appendix A

Organization of the I BP in the United States Richard C. Dugdale (Biological Productivity in Upwelling Eco- systems) Stanley P. Gessel (Coniferous Forest Biome) More than 3 years of planning and coordinating David W. Goodall (Desert Biome) activities and 1 year of research sponsored by the U.S. Harold Mooney (Mediterranean Scrub Project) Dieter Mueller-Dombois (Island Ecosystem Stability and National Committee have culminated in a sophisticated Evolution Subprogam) scientific plan of integrated and coordinated research Howard T. Odum (Tropical Forest Biome) programs. The IBP organization now consists of: G. Carleton Ray (Marine Mammals) Otto T. Solbrig (Structure of Ecosystems Subprogram) Executive Committee -George M. Van Dyne (Grasslands Biome) Directors of integrated research programs Directors of coordinated research programs Directors of biomes Human Adaptability Component Directors of key subprograms Paul T. Baker (Biology of Human Populations at High Altitudes) This membership constitutes the U.S. National Com- William S. Laughlin (International Study of Circumpolar mittee for the IBP. Peoples) Frederick A. Milan (International Study of Eskimos Sub- program) James V. Neel (Population Genetics of South American Indians)

EXECUTIVE COMMITTEE W. Frank Blair, Chairman COORDINATED RESEARCH PROGRAMS T. C. Byerly, Vice-Chairman Stanley I. Auerbach. Vice-Chairman Paul T. Baker Environmental Component Everett S. Lee Frederick Sargent, II William S. Benninghoff (Aerobiology) Frederic H. Wavier John L. Creech (Conservation of Plant Genetic Materials) Donald S. Farner (ex officio) C. C. Delwiche (Biology and Ecology of Nitrogen) Carl B. Huffaker (Biological Control of Insect Pests) George Sprugel, Jr. (Conservation of Ecosystems) Forest Stearns (Phenology) INTEGRATED RESEARCH PROGRAMS

Environmental Component Human Adaptability Component

Stanley I. Auerbach (Deciduous Forest Biome) C. Glen King (Nutritional Adaptation to the Environment) W. Frank Blair (Origin and Structure of Ecosystems) Everett S. Lee (Biosocial Adaptafion of Migrant and Urban Jerry Brown (Tundra Biome) Populations)

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118 Appendix B

Officers and Members of the Special Committee Convenors of sectional committees for the International Biological Program PT Prof. J. B. Cragg (Canada) PP Academician Malek (Czechoslovakia) CT Mr. E. M. Nicholson (United Kingdom) PF Prof. G. G. Winberg (Union of Soviet Socialist OFFICERS Republics) PM Prof. M. Dunbar (Canada) HA Prof. J. S. Weiner (United Kingdom) President UM Dr. G. K. Davis (United States)

Prof. F. Bourliere (France) Representing internatiorI scientific unions other than those in ICSU Vice-Presiden ts IUCN Prof. J. G. Baer (Switzerland) Prof. W. F. Blair (United States) IUAES Prof. J. Hiernaux (Belgium) Sir Otto Frankel ;Australia) IAHB Prof. J. Hiernaux (Belgium) Academician I. Mdlek (Czechoslovakia) Prof. H. Tamiya (Japan) Representing scientific committees of ICSU

Scientific Director SCAR Prof. G. A. Knox (New Zealand) SCOR Dr. 0. H. Oren (Israel) Dr. E. B. Worthington (United Kingdom) Elected members

Prof. W. F. Blair (United States) Dr. A. E. Boyo (Nigeria) MEMBERS Prof. B. E. Bychowsky (Union of Soviet Socialist Republics) Prof. F. di Castri (Chile) Sir Otto Frankel (Australia) Representing ICSU and member unions of ICSU Dr. R. W. J. Keay (United Kingdom) ICSU Prof. D. Blaskovic (Czechoslovakia) Prof. G. Montalenti (Italy) IUBS Dr. Arthur D. Hasler (United States) Prof. K. Petrusewicz (Poland) I UB Prof. M. Florkin (Belgium) Dr. F. Salzano (Brazil) I UPS Prof. P. 0. Astrand (Sweden) Prof. B. R. Seshachar (India) IGU Prof. C. Troll (Federal Republic of Germany) Dr. H. Tamiya (Japan) I UPAB Dr. A. R. Gopal-Ayengar (India) Prof. H. Thamdrup (Denmark) IUNS Prof. C. G. King (United States) Prof. C. A. du Toit (Republic of South Africa)

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Countries Participating in the I BP, with Chairmen Indonesia Dr. 0. Soemarwoto Ireland, Republic of Mr. F. McDonnell of National Committees Israel Prof. M. Evenari Italy Prof. C. Barigozzi Prof. H. Tamiya Argentina Prof. 0. Boelcke Japan Australia Sir Otto Frankel Korea, Republic of Prof. Yung Sun Kang Dr. N. P. Mwanza Austria Prof. W. Kiihnelt Malawi Enche K. D. Menon Belgium Prof. P. Duvigneaud Malaysia Dr. A. Ayala-Castaiiares Brazil Prof. A. Cury Mexico Prof. G. J. Verve 1de Bulgaria Prof. I. Pashev Netherlands Prof. G. A. Knox Canada Dr. T. W. M. Cameron New Zealand Prof. C. I. 0. Olaniyan Ceylon Prof. B. A. Abeywickrama Nigeria Prof. R. Vik Chile Prof. Dr. F. di Castri Norway China, Repu Ltic of Dr. Jong-Ching Su Panama Dr. 0. Sousa Prof. Hernando de Macedo Colombia Dr. Luis Eduardo Mora Peru Dr. S. M. Cendafia Congo. Democratic Mr. J. Ileo Philippines Republic of the Poland Prof. K. Petrusewicz Prof. E. Bursell Czechoslovakia Prof. I. NM& Rhodesia Denmark Prof. H. M. Thamdrup Romania Prof. N. Salageanu Prof. C. A. du Toit East Africa Prof. W. B. Banage South Africa, Kenya Dr. R. S. Odingo Republic of Tanzania Mr. A. S. Msangi Spain Prof. A. Carrato Uganda Dr. G. H. Kiwauwa Sweden Prof. C. 0. Tamm Ecuador Dr. M. Acosta-Solis Thailand Lt. Gen. P. Salvidhannidhes Finland Prof. H. Luther Tunisia Mr. El Fekkih France Prof. Th. Monod Union of Soviet Socialist Prof. B. E. Bychowsky Germany, Democratic Prof. H. Stubbe Republics Republic of United Arab Republic Prof. H. Said Germany, Federal Prof. H. Ellenberg United Kingdom Prof. A. R. Clapham Republic of United States Prof. W. F. Blair Dr. C. Estable Ghana Dr. A. J. E. Bucknor Uruguay Prof. F. H. Weibezahn Greece Prof. G. Athanasiades Novas Venezuela Hungary Prof. J. Balogh Vietnam, Republic of Mr. Pham-Hoang-Ho India Prof. B. R. Seshachar Yugoslavia Prof. Dj. Jelenic

121 120 fe.t4