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University Micrdnlms International 300 N ZEE B ROAD, ANN AR80R. Ml 48106 18 BEDFORD ROW. LONDON WC1R 4EJ. ENGLAND 8100158

G o l d m a n , J a c k H e r b e r t

COMPARATIVE DIETS AND REPRODUCTIVE OUTPUT IN AN OMNIVOROUS INSECT

The Ohio State University Ph.D. 1980

University Microfilms International300 N. Zeeb Road, Ann Aibor, MI 48106 COMPARATIVE DIETS AND REPRODUCTIVE OUTPUT IN AN OMNIVOROUS INSECT

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By Jack Herbert Goldman, B.S., M.S.

The Ohio State University 1980

Reading Committee* Approved by Dr. George Dalrymple Dr. David Denlinger Dr. David Horn Environmental Biology Program Dr. Sheldon Lustick To my wife and parents, who made this worth doing. ACKNOWLEDGMENTS

I would like to thank George Dalrymple for his faith in me, his patience, his inexhaustable good will, and his invaluable assistance! there are few people like George. I also want to thank my wife, Jan, for her tireless interest and support and help in writing and the graphics* I promise, Jani never again! I also want to thank Dave Denlinger for his help and the use of his lab, and also Shelley Lustick, Dave Horn, Roy Tassava, Pete Pappas, George Barber, Rodger Mitchell, Mike Murtaugh, Ann Gnagey, Jay Bradfield, John

McCabe, Sue Mahaney, Ron Stuckey, the Environmental Biology Program, and the Department of Zoology. VITA

July 15» 1 9 5 ^ .... Born, New York City 1971-1975 ...... New York State Regents Scholarship 1975 ...... Teaching Assistant, State Univer­ sity of New York at Binghamton

1975 ...... B.S., Biological Sciences, State University of New York at Binghamton

1975-1979 ...... Graduate Fellow, The Ohio State University, Columbus, Ohio 1976-1980 ...... Graduate Teaching Associate, The Ohio State University, Columbus, Ohio 1978 ...... M.S., Environmental Biology, The Ohio State University, Columbus, Ohio

FIELDS OF STUDY Nutritional physiology of cockroaches Computer simulation of population extinction Theoretical ecology TABLE OP CONTENTS Page DEDICATION ...... ii ACKNOWLEDGMENTS ...... iii

VITA ...... iv LIST OF T A B L E S ...... vi LIST OF FIGURES...... ix Chapter I. INTRODUCTION ...... 1

II. MATERIALS AND METHODS ...... 6

A. Organization of Study ...... 6 B. Husbandry...... 6 C. Sampling ...... 9 D. Measurements ...... 10 E. Calculations...... 11 F. Statistical analysis ...... 16 III. RE S U L T S...... 18 A. Growth and demography...... 18 B. Food c o n s u m p t i o n ...... 21 C. Approximate digestibility ...... 21 D. Body w e i g h t ...... 22 E. Relative growth r a t e ...... 23 F. Conversion efficiencies ...... 23 G. Oxygen consumption ...... 2k H. Reproductive investment ...... 2k I. Composite food u s e ...... 25 IV. DISCUSSION ...... 26 V. CONCLUSIONS...... kO

BIBLIOGRAPHY ...... kk

v LIST OF TABLES

Table Page 1. Pooled number of survivors at each instar for each diet (juvenile period), and G-test for differences (if significant, diets listed in increasing order by initial* L, low pro­ tein; M, medium protein; H, high protein) .... 58

2. Days to adulthood, each d i e t ...... 58

3 . Pooled number of female survivors at begin­ ning of adulthood and at first three repro­ ductive efforts for each diet, and G-test for differences...... 59 4. Interval (days) between adult eclosion of females and the appearance of the first ootheca...... 60 5. Pooled number of offspring produced at first three reproductive efforts for each diet, and G-test for differences...... 6 l

6 . Pooled values of number of offspring produced per reproductive effort at first three repro­ ductive efforts for each d i e t ...... 61 7. Pooled values of percentage of successful reproductive efforts at first three repro­ ductive efforts for each diet ...... 62

8 . Pooled values of number of offspring produced per successful reproductive effort at first three reproductive efforts for each diet...... 62 9. Medians and ranges (in parenthesis) of values of approximate digestibilty ...... 63 10. Medians and ranges (in parentheses) of the consumption index for each diet at each sampling period, expressed as mg consumed per mg body weight per day (all weights are dry weights). . . 66

vi LIST OP TABLES (continued)

Table Page 11. Medians and ranges (in parentheses) of values of body weight (in mg dry weight) for each diet at each sampling period (ten individuals sampled per d i e t ) ...... 68 12. Pooled values of relative growth rate (in mg gained per mg weight (dry weights)), each sampling period, for each d i e t ...... 70 13. Pooled values of conversion efficiency of ingested food (as percentages, based on dry weights) for each d i e t ...... 71 14. Pooled values of conversion efficiency of (digested) food into body substance (as per­ centages, based on dry weights) for each diet . 72 15. Medians and ranges (in parentheses) of oxygen consumption (expressed as microliters 0 2 per mg dry weight per day) for each diet at each sampling (one individual sampled per d i e t ) ...... 73 16. Pooled values of reproductive investment (expressed as total mg. dry weight of oothecae produced per mg dry weight of Insect per day) for each d i e t ...... 75 17. Reproductive biomass produced per day as a percentage of total food ingested per day (dry weights), each sampling period, for each d i e t ...... 76 18. Reproductive biomass produced per day as a percentage of toal food digested per day (dry weights), each sampling period, for each d i e t ...... 76 19. Medians and ranges (in parentheses) of dry weights of second-effort oothecae (in mg.) for each d i e t ...... 77 20. Pooled values of number of oothecae per female per day for each population sampling period, each d i e t ...... 7 7

vii LIST OF TABLES (continued)

Table Page

21. Food use values of select insect species (other indices* CIt consumption index| R, respiration + ingestions RA, respiration assimilation)...... 78 22. Averages and standard deviations of food use values of selected insect orders from Table 21 and Blattella germanica ...... 82 23* Compensations in food use (+, positive evi­ dence for mechanism* negative evidence for mechanism)...... 83 24. Age or size-related trends in food use indices (+t increase* -t decrease* 0 , unclear) .... 86

viii LIST OP FIGURES

Page Figure 1. Fooled number of adult survivors in popula­ tions fed each diet through the first three reproductive efforts. (Solid line, high protein diet* dashed line, medium protein diet; dotted line, low protein diet ) . . . 90 2. Fooled number of offspring from populations fed each diet through the first three repro­ ductive efforts. (Solid line, high protein dieti dashed line, medium protein diet; dotted line, low protein diet ) ...... 91

3 - Percentage of reproductive efforts, pooled for populations fed each diet, that yield offspring, through the first three repro­ ductive efforts. (Solid line, high protein diet) dashed line, medium protein diet; dotted line, low protein diet ) ...... 92

Medians of pooled values of approximate digestibility (expressed as a percent) at each sampling for populations fed each diet. (Cross-hatched areas, high protein diet; dotted areas, medium protein diet; solid white areas, low protein diet ) ...... 93

5. Composite of the fate of ingested food (i.e., food use) during the lifetime of individuals fed the high protein diet...... 6. Composite of the fate of ingested food (i.e., food use) during the lifetime of individuals fed the medium protein diet...... 95

7. Composite of the fate of ingested food (i.e., food use) during the lifetime of individuals fed the low protein diet...... 96

ix CHAPTER I

INTRODUCTION

The present study complements previous work on the quantitative nutrition of insects. The food use and repro­ ductive output of Blattella germanica L. (Dictyopterai Blattellidae), the German cockroach, were monitored for its entire lifespan, using a control diet and two other diets diluted with dextrose. All three diets contained protein levels in the "optimal** range of B^ germanica (Haydack,

1953)1 so that the compensations made would reflect normal homeostasis rather than stress-induced responses. Differ­ ences in performance within the optimal range would also "be seen. This work provides a view of physiological adapta­ tion to protein depletion by a scavenger species, and thus supplements previous studies done on the processing of food by insects. Nutritional studies on insects emphasize energy and raw materials needed for growth, homeostasis, and reproduction

(House, 1969). Qualitative studies determine the substances an insect must extract from its environment because these substances cannot be synthesized (Fraenkel, 195° 1 Sang,

19561 Gordon, 1959* c.f. House, 197*0* In contrast, quan­ titative studies examine the proportions of required 1 z nutrients necessary for the insect to function normally

(Waldbauer, 19681 House, 1969)• Rates of growth, development, and mortality have com­ monly been used as the initial criteria in studies of diet in insects (e.g., Haydack, 1953* Atwal, 1955* Gordon, 1968). An insect can use two diets differently to produce the same demographic patterns (House, 1966* Soo Hoo and Fraenkel,

1966). In order to distinguish between diets, the quantity of food eaten, its digestibility, and the efficiency of con­ version of food into biomass are also considered in quanti­ tative studies (Waldbauer, 1968* Waldbauer and Bhattacharya, 1973). Early work on food use included studies on the silk moth, Bombyx mori (Hiratsuka, 1920, in Waldbauer, 1968), the cabbage aphis, Brevicorvne brassicae (Evans, 1938), and the desert locust, Schistocerca gregaria (Davey, 195*0* Most quantitative studies deal with phytophagous (leaf-eating) insects, due in part to their predominance

(Fraenkel, 1959* House, 1969) and their economic importance. Recent examples include studies on the Mexican bean beetle,

Eoilachna varivestis (Barney and Rock, 1975)* "two species of grasshoppers, Melanoplus spp. (Bailey and Mukerji, 1976), and the gypsy moth, Lymantria dispar (Barbosa and Greenblatt,

1979). These studies often examine the effect of allelo- chemic or secondary plant substances on insect feeding be­ havior and food use (McGinnis and Kasting, 1966* Soo Hoo and 3

Fraenkel, 1966* Latheef and Harcourt, 1972* Hoekstra and Beenakkers, 1976) Reese and Beck, 1976a,b) Dahlman, 1977) Scriber, 1978a,b, 1979a,bj Scriber and Feeny, 1979)• The comparison of specialist and generalist feeders generates discussion on the metabolic costs of maintaining detoxifi­ cation systems (Fraenkel, 1959) Feeny, 1976* Schroeder,

1976) and Scriber, 1977)1 and ultimately on the power ver­ sus efficiency tradeoffs of Odum and Pinkerton (1955) (Me­ drano and Gall, 1976) Schroeder, 1977* Slansky and Feeny,

1977) Scriber, 1978a). There are two important features of food use noted in previous work* 1) to maintain relative nutrient intake,

insects increase their consumption when a diet is diluted with a sugar or cellulose (Dadd, I960* House, 1965) McGin­ nis and Kasting, 1967) Bignell, 1978)* 2) to produce a con­ stant gross conversion efficiency (percentage of ingested food converted to biomass), assimilation efficiency and metabolic rate vary inversely. Researchers have used this difference in food use to explain contrasting assimilation efficiencies and metabolic rates between herbivores and car­

nivores (Welch, 1968) Calow, 1977)* Herbivorous insects fed different plants have also demonstrated the second fea­

ture. Examples include selected lepidopterans (Evans, 1939)» the Colorado potato beetle, Leptinotarsa decemlineata (La­ theef and Harcourt, 1972), Tribolium (Medrano and Gall, 4

1976), and the black cutworm, Agrotis ipsilon (Reese, 1978). Few studies of insects examine non-herbivoresj the

German cockroach is a scavenger and an omnivore. It eats solid food, eliminating the problems of handling live plant material (see Waldbauer, 196**, 19681 Gordon, 1968) and plant allelochemics. Diet manipulation is also easier. Me­ tabolic pathways in cockroaches are different from other in­ sects, due to the presence of bacterial symbiotes in the fat body (Brooks and Richards, 1955* Gordon, 1959; Henry, 1962). Monitoring food use and reproductive effort in germanica also differs from previous work because the insect is hemi- metabolous and iteroparous. Grasshoppers and locusts, which are also hemimetabolous and iteroparous, have been examined

(e.g., Clarke, 195?! Smalley, i9 6 0j Mordue (Luntz) and Hill, 1970j and Mispagel, 1978), but they are phytophagous, and have nutritional requirements different from blatellids.

The present study examined the effects of decreasing protein intake in the German cockroach by 50%. germani- ca ingested the same amount of food, converted it into bio­ mass with the same efficiency, and invested the same propor­ tion of food into reproduction, producing an equal number of offspring per ootheca (egg case). The decreased protein intake did result in higher mortality, slower growth rate, and smaller individuals, but these differences are minimal in relation to the change in protein intake. Such 5

flexibility in germanica is best explained by its ability

to recycle nitrogenous wastes (urates) to supply energy and nitrogenous compounds, with the help of bacterial symbiotes located in the fat body. CHAPTER II

MATERIALS AND METHODS

A. Organization of study Populations of Blattella germanica (derived from En­ tomology Department stocks) were raised on three diets.

The first diet was Purina Lab Meal (Ralston-Purina Co.), which contained 2 3 .5 # protein by weight ("high protein")* the second and third were diluted with dextrose by 25# and

50 # to yield protein contents of 17 .6# ("medium protein") and 11.8# ("low protein").

Twenty populations were raised on each diet. Ten of the populations were experimental populations that were sam­ pled during the study. The other ten provided replacements of the same age at each sampling which were raised under i- dentical conditions. Each population was started when five oothecae from stock populations hatched in a 2^-hour period, providing an average of 139 individuals.

B. Husbandry

All populations were kept in an isolated room with no windows. Photoperiod was 12 hours lighti 12 hours dark.

The temperature was maintained at 27.il- + 1.1°C with an elec­ tric fluid-filled baseboard heater (Hydronic, model EP-1500, 5100 Btu/hr). Relative humidity was kept at 35.0 + 1.7# with a portable dehumidifier (Westinghouse, model ED-2078, 20 pints/day capacity). The low relative humidity preven­ ted food spoilage. All stages of each population were housed in 32 x 154 x 10 cm clear plastic boxes. Fewkes (i9 6 0) reported a greater metabolic cost due to searching for food by smaller instars of the nabadid bug, Stalia major, kept in the same size box as adults. His conclusions ignore a greater meta­ bolic rate in younger instars, as found for the dusky grass­ hopper, Encoptolophus sordidus costalis (Bailey and Riegert, 1972), the blowfly, Lucilia illustris (Hanski, 1976), the rice weevil, Sitophilus orvzae (Singh, Campbell, and Sinha, 1976), the mite Caloglvphus berlesei (Stepien and Rodriguez, 1972), and of particular interest, Bj_ germanica (Woodland,

Hall, and Calder, 1968). Containers were therefore not scaled to instar size in the present study, since most evi­ dence points to a normally-occurring decrease in metabolic rate with age. A rectangular hole (18 x 11 cm) was cut in each opaque plastic lid and fitted with 32 x 32 nylon mesh.

Shelters were supplied to each container to provide protec­ tion for molting nymphs.

Food was prepared in pellet form on a Farr pellet press, using a mixture of the food compound and water. The pellets were 1.4— 1.7 cm high, with a diameter of 1 .3 cm. All pel­ lets were dried and stored at 60°C. Water was provided in

1 dram (15 x 45 mm) vials stoppered with 74 cm cotton dental wicks. Both food and water were supplied ad libitum. Each experimental population was monitored at least

once every three days. Food and water levels were checked, and a census taken of the population. Survival and devel­

opmental rates in nymphal populations were recorded for males and females, since in the first five instars, sexes can only be distinguished by microscopic examination (see

Ross and Cochran, i960). An average population instar was calculated at each census; when the average instar ap­ proached an integer, the population was sampled.

Adult females were easily recognized and individually marked on their wings at eclosion. A system of enamel paint

dots of different colors was used to distinguish females from each other. Such marks would not be lost because un­ like Periplaneta americana. the American cockroach, wings of germanica do not become damaged or removed due to agonistic behavior (personal observations). Adult female emergence and survival were recorded at

each census. Oothecae and offspring produced by individual females were also recordedt newly-hatched offspring were al­ ways removed. An adult population was sampled when, on the average (at the population level), the first (adult sampling "A")» second (adult sampling "B"), or fourth (adult sampling

"C") ootheca ("reproductive effort") was produced. C. Sampling Growth, demography, and food use data of juvenile populations are organized around the sampling--the integral average instar. Adult survival and reproductive data are grouped around adult emergence and reproductive efforts, since each individual female *s performance could be monitor­ ed. Only data for the first three reproductive efforts are used, since at least of all offspring had been produced by the third effort (see Table 4a)j beyond the third repro­ ductive effort, sample sizes dropped and sampling error tended to mask differences. Adult food use data (e.g., food consumption, faeces production) was based on population performance. Asynchrony of reproductive efforts necessita­ ted collection of the data at sampling times based on the average reproductive effort.

Each population was anaesthetized with carbon dioxide before sampling. Brooks (1957t 1965) found that regular carbon dioxide application retarded growth rates, decreased adult weights, reduced the number of offspring, and pro­ longed adult life; other gases tested (e.g., nitrogen, nitrous oxide, ethyl ether) had more pronounced effects.

Gordon (1972) has disputed her findings, claiming carbon dioxide only affected one growth stage. The consequences of anaesthesia were not considered important, since in any event carbon dioxide was used uniformly on all populations, regardless of diet. 10 Four individuals were removed from each sampled juvenile population! the remaining population was moved to a new container. A female, sexed after Ross and Cochran

(i960), was selected from the four individuals. The other three were returned to the population, along with a female from the associated replacement population. Adult popula­ tions were sampled by removing a female and replacing it with a female from the replacement population. The substi­ tute was marked with red enamel paint to distinguish it from the other females, to prevent her from being subse­ quently sampled. The food pellets were gathered from the emptied con­ tainers and stored at 60°C. New pellets of known dry weight were supplied to the new container. Faeces were collected and frozen. Females were frozen after oxygen consumption was measured (see below)} oothecae from adult females of sampling B were removed before respirometry and also stored at 0°C.

D. Measurements

The difference in food pellet weights before and after feeding was used to determine ingestion rates. All pellets were stored at 6o°C for at least ten daysr they were then weighed to constant weight on a Mettler PI63 balance. B_. germanica is considered to be a clean eater and does not leave faeces on its food (Brooks, 1965). 11

Oxygen consumption of sampled females was determined in a microvolumetric respirometer (Scholander, Claff, An­ drews, and Wallach, 1952). The water bath was maintained at 25°C; readings were taken over a 4-24 hour period, de­ pending on the size (and respiratory rate) of the individual. Expired carbon dioxide was absorbed by 10# potassium hy­ droxide . Faeces, oothecae, and sampled individuals were stored at 0°C or less for up to ten days in preparation for freeze- drying. The samples were then lyophilized in a Virtis Model 2040 Freeze-dryer for 18 hours, and stored for up to four days in a desiccator. The samples were then weighed on a Cahn Gram Electrobalance to the nearest 0.001 mg.

E. Calculations

Waldbauer*s (1964, 1968) indices of food use were used:

consumption index - _____ weight of food______, 1. time (days) x weight of animal 1 '

relative growth rate = _____ weight gain during feeding period______time (days) x weight of animal (beginning period) ' ^

efficiency of conversion of ingested food to body substance (gross conversion efficiency) = relative growth rate inn consumption index 12

approximate digestibility = (weight of food ingested) - (weight of faeces) 1nn (weight of food ingested)

efficiency with which (digested) food is converted to body substance (net conversion efficiency) - ______(weight gained)______100 (weight of ingested food) - (weight of faeces)

Metabolic rate can be calculated from the following equation (Waldbauer, 1968)1

metabolism = consumption - faeces - growth (6), where all terms, including metabolism, are expressed as bio­ mass (dry weight). A direct measurement of oxygen consump­ tion, however, was used in the present study to determine metabolic rate. Several studies have shown that the Wald­ bauer (1968), or gravimetric method, yields consistently higher values than the oxygen consumption method. Schroeder

(1972, 1973) found that gravimetric values were 1.7 times those of oxygen consumption in the cecropia moth, Hvalopho- ra cecropia. and 1 .5 in the moth, Pach.vsphinx modestat Wood­ land, et al. (1968) found gravimetric determinations to be 5# greater than oxygen consumption values in third instars of germanica. and 20# greater in adults. The inherent variability of oxygen consumption measurements is also great. Standard metabolism is augmented by energy used for movement and digestion in the insect is not starved and is free to 13 move (Sweeney and Schnack, 1977)* The ratio of insect vol­ ume to specimen tube volume will decrease as the animal grows. Clarke (1957) changed specimen tubes in his Warburg apparatus so that no insect was greater than 5# of the tube volume. In the present study, oxygen consumption measure­ ments were used because it is a more direct measurement of metabolism* the gravimetric method is subject to error from a variety of sources not directly related to metabolic rate. Unstarved and active animals were used in standard-size tubes for oxygen consumption measurements to keep the pro­ cedure simple* such potential sources of error are constant for all three diet treatments. The biomass lost as exuviae is usually not added to production to get "true" production figures (Waldbauer, 1968). This can be a considerable source of error. Several

studies have estimated exuviae loss* 0 .1# of net production

in the blowfly, Lucilia illustris (Hanski, 1976), 4# in the paddy field grasshopper, Oxva velox (Delvi and Pandian,

1971)i 5 *15 $ in the chrysomelid beetle, Melasoma collaris

(Hagvar, 1975). 5-7# in the grasshopper, Bootettix punctatus (Mispagel, 1978). 6-1155 in the dusky grasshopper, Encopto- lophus sordidus costalis (Bailey and Riegert, 1972), and

7-11# in the moth, Cvclophragma leucosticta (Mackey, 1976). Many insects, including the pea aphid, Ac.vrtho siphon pi sum

(Randolph, Randolph, and Barlow, 1975). the range caterpil­ lar, Hemileuca oliviae (Schowalter, Whitford, and Turner, 14

1977)» and most importantly, the German cockroach (Woodland, et al., 1968) consume their exuviae after molting. The re­ cycling of that biomass reduces that potential error, and no correction has beeh made for exuviae. The measurement of faeces produced also involves error. Waldbauer (1968) noted that Bi germanica probably eats its faeces, although he provides no estimate of how much is consumed; consumption would tend to lower the gross conversion efficiency (Waldbauer, 1968). Faeces consumption is not mentioned in an energy budget study of the roach

(Woodland, et al., 1968). Faeces also contain excreted peritrophic membranes and digestive enzymes from the gut

(Woodland, et al., 1968; Schroeder, 1976). These consti­ tuents of the faeces have also not been quantified. No cor­ rection has been made for either error as a result.

Retention of food in the gut will not be corrected for in the present study, although it is a source of error

(Waldbauer, 1968). Food was found to make up 9*3# "the fresh body weight of the tobacco hornworm, Manduca sexta (Waldbauer, 1962) and 1.2# in the migratory locust, Locusta migratoria (Clarke, 1957); this figure was 4.4# in ger­ manica (McCay, 1938). Retained food causes an overestima­ tion of approximate digestibility, as well as conversion efficiencies and biomass. The error is consistent in the present study, and is not considered significant. 15

A basic problem affecting studies on insects other than B_j_ germanica is the excretion of faeces combined with metabolic nitrogenous wastes, usually as uric acid (Wald- bauer, 1968* Bhattacharya and Waldbauer, 1972i Gordon, 1972). Approximate digestibility was increased less than 1% in Bombyx mori when uric acid was subtracted from the faeces (Hiratsuka, 1920, in Waldbauer, 1968)1 other correc­ tions for uric acid resulted in an increase in approximate digestibility of k.75% in the mealworm, Tenebrio molitor (Evans and Goodcliffe, 1939)# and an increase of 3.7“7*0# in larval Tribolium confusum (Bhattacharya and Waldbauer,

1972). Correcting for uric acid also slightly decreases the efficiency of converting digested food. The cockroach is different since it stores virtually all of its uric acid in the fat body (only 2$ of it is excreted). The uric acid is used in subsequent metabolism (Mullins and Cochran, 1975a,b); as a result, production is overestimated by only 1% (Gordon, 1959)* Ammonia is the primary nitrogenous ex­ cretory product in B.*. germanica (2k% in females* Mullins and

Cochran, 1972, 1973# 197^# 1975a). Much of the ammonia (52%) is lost as the faeces dry (Mullins and Cochran, 1973), so less nitrogenous waste is found in the excreta than in other insects. Estimates of approximate digestibility based on faeces mass should be more accurate as a result. 16

F. Statistical analysis Since most sample sizes were ten or less, nonpara- metric statistical procedures were used to analyze the data

(Hollander and Wolfe, 1973)- The primary methods used were tests to detect differences between several samples with replicates (one-way layouts: Kruskal-Wallis and Jonckheere-

Terpstra tests, Hollander and Wolfe, 1973) an

The G-test (Sokal and Rohlf, 1969) was used to compare fre­ quency data.

The statistic calculated in the Friedman test, Kruskal-

Wallis test, and G-test is compared to a critical value from the X distribution (Sokal and Rohlf, 19691 Hollander and Wolfe, 1973)• The notation of the critical value p in the text is, for example, x ( 05 2) “ 5 *9 9 1* where .05 is the significance level (5 $)* and 2 is the number of de­ grees of freedom. The Page test statistics are compared to valued of a distribution described by Hollander and Wolfe (1973*372). An example of the critical value notation is

1 (,0 5 ,3 ,5 ) = w*iere *°5 significance level, 3 is 17 the number of samples, and 5 the number of replicates. The J* statistic of the Jonckheere-Terpstra test is compared to the 5% cutoff from the normal distribution. Sample sizes are designated when this test is used. Significance levels are symbolized in the tables next to the test statistic* *, significant at the 5# level; **, significant at the 1# level; ***, significant at the 0.5# level. The symbol "NS" denotes no significance at the 5% level. CHAPTER III RESULTS

A. Growth and demography Data were pooled for all populations fed the same diet because of high variability due to small sample sizes*

1. Juveniles There were no overall differences associated with diet in juvenile survivorship (males and femalest see Table 1). No consistent differences due to diet existed at hatching (G-test, G = 3*5^8, gj = 5*991. P > *05, Sokal and Rohlf, 1969*560) or through the first four instars Survival in the populations fed the low protein diet was lower at instars 5, 6, and adult eclosion than in popula­ tions fed the medium and high protein diets (Table It G-test P < .05. Sokal and Rohlf, 1969*560). Differences due to diet also existed in the develop­ mental rate of individuals of the populations (Table 2). Individuals of populations fed the medium protein diet developed fastest, followed by populations fed the high protein diet, and then those fed the low protein diet

(Jonckheere-Terpstra test, J* = 1*7^9. = 1*6^5.

P < .05. n = 10, Hollander and Wolfe, 1973*120). 18 19

2. Adults Adult female survival continued the trend started at the end of the larval stage. The populations fed the low protein diet had the lowest survival at all four samplings; populations fed the medium protein diet always had the highest survival (Table 3a; Figure 1). Survival of popula­ tions fed the medium protein diet was always significantly higher than that of populations fed the low and high protein diets, based on paired comparisons at each sampling using the G-test (Table 3b; P< .005. Sokal and Rohlf, 1969*560). There were no diet-related differences in the number of days between adult eclosion and the first ootheca pro­ duced (Table 4; Kruskal-Wallis test, H = 1.644, X2^ -

5-991, P> .05, Hollander and Wolfe, 1973*115)* Differences in reproductive output associated with diet reflect the differences seen in survival (Table 5a; Figure 2). Popula­ tions fed the medium protein diet always produced the most offspring (9062 total), and populations fed the low protein diet always produced the fewest (5748; populations fed the high protein diet produced 6413 offspring). Paired compari­ sons using the G-test (Sokal and Rohlf, 1969*560 ) indicate that these differences are always highly significant (P< .005; Table 5b). The same highly significant differ­ ences existed in lifetime reproductive output (G-test,

G = 858 .6 , x2(t05,2) = 5*991. P<*005, Sokal and Rohlf, 1969*560). 20 A highly significant diet-associated trend existed in the number of offspring produced per ootheca (Table 6).

The populations fed the low protein diet yielded the most offspring per ootheca (or effort* 22.992 total), followed by populations fed the medium protein diet (21*942), and then by populations fed the high protein diet (18.6971 Page test,

L = 5 6 , 1^ 3 4 ,) ~ 5^» P < *005, Hollander and Wolfe, 1973*147)• Data on the number of offspring produced per ootheca can be partitioned into the percentage of successful oothecae produced and the number of offspring per successful ootheca. The populations fed the low protein diet had the highest percentage of successful oothecae (7 8 . total), followed by populations fed the medium protein diet (6 9*776 )» and then by the populations fed the high protein diet

(6 5 .O%); these differences are significant (Table 7»

Figure 3* Page test, L = 42, 1^ 05 3 3) = ^1| P < *01* Hollander and Wolfe, 1973tl47). No diet-related differences were seen in the number of offspring yielded per success­ ful ootheca (Table 8). Although the populations fed the low protein diets were more efficient in producing young than the populations fed medium and high protein diet, the advantage is offset by lower survival. As a result, the populations fed the medium protein diet produce the most offspring, and the 21 populations fed the low protein diet produce the fewest (Table 5a).

B. Food consumption There were no consistent diet-related differences seen in the consumption index (food ingested per mg dry body weight* equation (1) in Materials and Methods) in either the juvenile or adult phases (Table 10a,b). The index decreased with increasing age in the juvenile instars (Table 10c* Page test, P < .005* Hollander and Wolfe, 1973* 14-7). The populations fed the medium and high protein diets continued this trend through adulthood (Table 10c* Page test, L = 133, 128, respectively, 1^ ^ * 128, P < .05, Hollander and Wolfe, 1973*14?), but low protein populations increased consumption (Table 10a,c* Page test, L = 132, 1{ ^ = 128, P < .05, Hollander and Wolfe, 1973tl4?).

C. Approximate digestibility Approximate digestibility values (the amount of ingested food not voided as faeces* see equation (4) in Materials and Methods) were highest at all samplings in the populations fed the low protein diet, followed by the popu­ lations fed the medium protein diet, and then the popula­ tions fed the high protein diet (Tables 9a,b* Figure 4*

Jonckheere-Terpstra test, P < .005, n = 10, Hollander and Wolfe, 1973*120). The values for the populations fed the medium and low protein diets were significantly greater than 22 the values of the populations fed the high protein diets throughout the larval stage (Table 9b; paired comparisons based on the Kruskal-Wallis test, P<.05, n - 10, Hollander and Wolfe, 1973*124). Approximate digestibility decreases significantly with increasing age in the juvenile stage

(Table 9c; Page test, P^.005» Hollander and Wolfe, 1973*147) and increases significantly during adulthood

(Table 9c; Page test, P<.01, Hollander and Wolfe, 1973*147).

D. Body weight There were no diet-related differences in dry body weight in the larval stage (Table 11a,b; Kruskal-Wallis test, P > .05, n = 10, Hollander and Wolfe, 1973*115)- Dif­ ferences associated with diet exist at the second and third adult samplings (Jonckheere-Terpstra test, J* = 4.487, 3.422, respectively, = 1.645, P <.005, n = 10, Hollander and Wolfe, 1973*120). Populations fed the medium and high pro­ tein diets had heavier individuals than populations fed low protein diets at adult sampling B (Table lib; paired comparisons based on the Kruskal-Wallis test, P < .0 5 , n = 10,

Hollander and Wolfe, 1973*124), but individuals of the pop­ ulations fed the high protein diet were heavier than those of the populations fed the other two diets at adult sampling

C (Table lib; paired comparisons based on the Kruskal-Wallis test, P<.05, n = 10, Hollander and Wolfe, 1973*124). In­ dividuals of the populations fed the low protein diet did 23 not significantly increase in weight during adulthood (Ta­ ble 11c; Page test, L = 124, ,0 5 ,3 ,10) = *°5, Hol­ lander and Wolfe, 1973*147).

E. Relative growth rate Differences associated with diet occurred in the rela­ tive growth rate (weight gain per mg initial wt.; equation (2) in Materials and Methods) during the larval instars. Individuals of populations fed the high protein diet had the highest rate of growth, followed by individuals of popula­ tions fed the medium protein diet, and then individuals of populations fed the low protein diet (Table 12; Page test,

L ■ 6 7 ,5 , 1^ ^ 5 ) = 66, P^.,05, Hollander and Wolfe, 1973* 14?). A decrease in growth rate with increasing age is also significant in all three diets (Page test, L = 157,

1 ( 05 5 3 ) = Hollander and Wolfe, 1973*1^7). There is a drop in relative growth rate in all diets from the sixth instar to adult sampling A.

F. Conversion efficiencies No diet-related differences were found in conversion efficiencies of ingested food into biomass (equation (3 ) in Materials and Methods) or of digested food into biomass

(equation (4) in Materials and Methods; Tables 13 and 14).

There were no differences associated with diet among juve­ niles (Friedman test, S = 2.800, X2(t05>2) = 5*991, P > .05, Hollander and Wolfe, 1973il40) or adults (Friedman test, 2Mr

S - 0 .6 6 7 , X2^ g) 5-991* P > .05, Hollander and Wolfe, 1973*1^0). No age-related trends were detected, except for the steep drop between instar 6 and adult sampling A.

Gr. Oxygen consumption There were no differences associated with diet in oxy­ gen consumption (per mg dry weight) in the juvenile stage

(Table 15a,bj Kruskal-Wallis test, all 2.681, X2^ 05 2) = 5.991, P>.05, Hollander and Wolfe, 1973*115)- Oxygen con­ sumption in the populations fed the high protein diet was generally lower in the adult stage than in the populations fed the other diets (Table 15b). Populations fed the medium diet had the highest oxygen consumption in the adult stage, followed by populations fed the low protein diet, and then populations fed the high protein diet (Page test, L = 41,

1( 05 3 3 ) “ *0^* Hollander and Wolfe, 1973*1^7)- Only the populations fed the high protein diet continued this trend into the adult stage (Page test, L = 130,

1 (-05,3,10) = 128, Hollander and Wolfe, 1973*1W •

H. Reproductive investment Reproductive investment is defined as the biomass in­ vested in oothecae per day per unit weight of insect. In the present study, there were no diet-related differences

(Table 1 6 ; Friedman test, S* = 1.151, X2^ 2) “ 5-991* P>.05, Hollander and Wolfe, 1973*1^0). Reproductive investment decreased with increasing age in adults (Page 25 test, L = 41, !(to5 3 3 ) = P ^ *05. Hollander and Wolfe, 1973il47). Data on reproductive investment can be partitioned into data on ootheca biomass and the number of oothecae produced per female per day. There were no differences associated with diet in the biomass of second-effort oothe­

cae (Table 19; Kruskal-Wallis test, H* = 3*259, X2^ Z ) s 5*991, P>.05, Hollander and Wolfe, 19?3ill6). Diet-re­ lated differences did not occur in the number of oothecae p produced (Table 20; Friedman test, S' = O.716 , X ^ 05 2) = 5.991, P/^.05, Hollander and Wolfe, 1973

1/v ne j , j ) = 41, P/.05, Hollander and Wolfe, 1973*147). There were also no diet-related differences in the pro­ portion of ingested biomass invested in reproduction (Table 17; Friedman test, S = 4.66 7 , X2^ Q^ 2j" 5-991, P^.05, Hol­ lander and Wolfe, 1973*140) and the proportion of digested biomass invested in reproduction (Table 18; Friedman test,

S = 4.667, x2(.o5 ,2) 5.991, P^.05, Hollander and Wolfe, 1973*140). Both proportions decreased with age (Tables 17 and 18; Page test, L = 41, 1^ ^ 3 ) = 41, P<.05, Hollan­ der and Wolfe, 1973*147).

I. Composite food use A composite of food use data for the lifespan of popu­ lations fed each diet is presented in Figures 5, 6 , and 7. CHAPTER IV DISCUSSION

There were several differences in growth, demography, and reproductive biology associated with diet in the present study of Blattella germanica. Protein content decreased by 50% by dilution with sugar from the high protein diet to the low protein diet. Approximate digestibility increased from an average of 73*85% in the populations fed the high protein diet to 84.86# in the populations fed the low pro­ tein diet (Table 9). These values are higher than in most insects (Tables 21,22), Approximate digestibility is high­ er for sugars than for proteins (Waldbauer, 1968; Beenak- kers, Meisen, and Scheres, 1971; Baker, 1975), so the amount of protein digested may be almost directly propor­ tional to the protein content of the food. Consumption rates did not differ significantly among populations fed the different diets (Table 10); these values were lower in B. germanica than in most insects (Tables 21,22). An adjustment in consumption and digestion to maintain the rate of assimilation, as reported in other insects (see Table 23), did not occur; the populations fed diets with different levels of protein probably digested approximately proportional amounts of protein. 26 Working with 5C$ less protein (and more dextrose)* the populations fed the low protein diet nevertheless had the same levels of consumption (see above), conversion (see Figures 5,6,7), reproductive investment, and individual reproductive output as populations fed higher levels of protein. There were diet-associated differences in juvenile and adult survival, growth rate, and weight of individuals* these differences are the product of a set of tradeoffs (see below) that enable the insect to function as normally as possible. Sugar intake was greater in diets with less pro­ tein* it is believed that protein shortage rather than sugar overabundance was critical in the present study (see below). The gross conversion efficiency may be the ultimate measure of food use, since it measures the amount of in­ gested food converted into biomass (equation (5) in Mater­ ials and Methods). The German cockroach did not alter the efficiency in response to a 50% change in protein intake (Table 13)* JLi germanica has a higher gross conversion efficiency than most insects (Tables 21,22)* values in pre­ vious studies on the German roach are 31.4?5 (McCay, 1938),

45% in an energy budget study (Woodland, et al., 1968), and

51-6^ (Brooks, 1965)- The Woodland, et al. (1968) figure may be high because energy budget studies often yield higher indices than biomass studies (see Schroeder 1971» 1973, 1976*

Van Hood and Dodson, 197*0* Brooks' (1965) value is suspect because she used wet weights, which may overestimate 28 conversion efficiencies by a factor of two to three (Fraen- kel and Blewett, 19^*M Waldbauer, 1968). Since the amount of protein assimilated is proportion­ al to protein intake, and no diet-related changes occurred in the gross conversion efficiency, it follows that there were no differences associated with diet in the net conver­ sion efficiency (Table 1*0, which is in the same range as other insects (Tables 21,22). Another means of measuring the net conversion efficiency is by monitoring the metabolic rate of an animal1 assimilated biomass is used either for production (conversion) or metabolism (Waldbauer, 1968). Differences in oxygen consumption would thus reflect dif­ ferences in the net conversion efficiency. No diet-related differences were found in juveniles in the present study

(Table 15). An adjustment has been observed in other insects be­ tween approximate digestibility and net conversion effi­ ciency to preserve the gross conversion efficiency (Table

23 ). No such compensation was observed in germanica. The only index of food use that exhibited diet-related dif­ ferences was approximate digestibility, and it probably did not result in increased protein assimilation. No other food use index changed in response to diet differences (see Figures 5»6,7)[ no compensations occurred among food use in­ dices to preserve either assimilation or conversion. Such a reaction points to the flexibility of germanica to 2 9 changes of as much as $0% in protein intake. On the level of processing the food, the animal did not make adjustments. Since it has less protein to work with, and did not de­ crease individual reproductive investment or output, the effects of less assimilated protein in the low protein diet must surface elsewhere. Individuals of 13^ germanica did not show diet-related differences in weight at any juvenile instar (Table 11). The decreased protein intake in populations fed the low pro­ tein diet resulted in a lower relative growth rate (Table 12) and a longer period of development (Table 2). Survival is also decreased in populations fed the low protein diet from the fifth instar through adult eclosion (Table 1).

The great sensitivity of juveniles to diet perturbations was discussed by Calow (1977). who spoke of a decrease in the ratio of "proliferative" to "functional" tissue as ani­ mals age. Juveniles, which have a high ratio, grow at a rate proportional to nutrient availability, since few stores exist and "building material" is at a premium. Ju­ veniles are also more sensitive to nutritional changes be­ cause the metabolic rate-conversion efficiency relationship (see above) is less flexible due to less plastic metabolic costs in younger animals (Calow, 1977). Slower growth and even decreased survival may thus be a result of poorer nu­ tritional quality, as in the present study. The changes in food use parameters as juveniles of B. germanica age and grow are similar to changes observed in other insects (Table 24). The consumption index de­ creases in all populations (Table 10), as does approximate digestibility (Table 9)• The relative growth rate (Table 12) and oxygen consumption (Table 15) also decline; the net conversion efficiency does not increase (Table 14) as in other insects (Table 24), and as predicted by a declin­ ing oxygen consumption rate. Gross conversion efficiency does not change as juveniles of B_;_ germanica (Table 13) or other insects (Table 24) develop. Diet-related differences in adulthood represent a set of tradeoffs probably mediated by the recycling of urates (see Mullins and Cochran, 1975a.b), with the help of bac­ terial symbiotes (Brooks and Richards, 1955; Valovage and Brooks, 1979)* The cost of decreased protein intake in the juvenile stage was slower development and increased mortal­ ity. The tradeoff in the adult stage is not as simple, since reproductive activity is superimposed over the in­ sect's other functions.

Adults fed the low protein diet had decreased survival

(Table 3 ), and lower weights with no weight gain (Table 11).

Fewer offspring were produced in populations fed the low protein diet (Table 5). but this only reflects decreased survival. The offspring produced per successful ootheca

(Table 8 ) and reproductive investment (Table 16) did not 31 reflect differences in diet. There was also a higher per­ centage of successful oothecae in the populations fed the low protein diet (Table 7)» possibly because the less hardy females fed the higher protein diets survived to adulthood, while those fed the low protein diet did not. The first eight to ten days after adult eclosion in B. germanica is a period of continued somatic growth, as in the desert locust, Schistocerca gregaria (Hill, Luntz, and Steele, 1968* Mordue (Luntz) and Hill, 1970* Walker, Hill, and Bailey, 1970)• During this period structures used by the adults (e.g., wings, ovaries) are developed (Mordue (Luntz) and Hill, 1970). Ingestion (Table 10) and metabol­ ic rate (Table 15) increase in B^ germanica during this period. The length of time between adult eclosion and the production of the first ootheca in the German cockroach does not show differences associated with diet (Table 4); in ad­ dition, all populations gained weight after adult eclosion (Tables 11 and 12). The protein to carbohydrate requirement ratio is 1.86 during somatic growth and 19*73 during subse­ quent ovarian development (in gregaria; Mordue (Luntz) and Hill, 1970), so protein depletion appears to be more critical after somatic growth is completed. Protein is a key component in reproductive production*

71# of dry oocyte weight comes from proteins in Sj. gregaria (Hill, et al., 1968). Protein levels during oogenesis have been examined in the grasshoppers gregaria (Hill, 1962) 32 and Locusta migratoria (Bar-Zev, Wajc, Cohen, Sapir, Apple- haum, and Emmerich, 1975)» and the cockroaches Nauphoeta cinerea (Rao and Fisk, 1965), Periplaneta americana (Mills,

Greenslade, and Couch, 1966; Bell, 1969)1 Diploptera punc­ tata (Stay and Clark, 1971). and Leucophora maderae (Wyss-

Huher and Luscher, 1972)1 levels increase to feed oogenesis and then decrease at the time of oviposition or ootheca pro­ duction. Tanaka (1973) and Tanaka and Ishiaki (197^) found similar protein behavior in B_^ germanica. Supplying protein for oogenesis can be crucial. Adult females of the desert locust, S^ gregaria. get most of the protein necessary for reproduction from ingested food

(Hill, et al,, 1968). Cockroaches are not as dependent on ingested food for protein as the desert locust (Donnellan and Kilby, 1967). But a constant food supply is still a ne­ cessity. Unfed females of germanica will not produce eggs; virgin females will not mate when starved (Roth and

Stay, 1962). Kunkel (1966) calculated that up to 90# of the food reserves of the German roach are used for each re­ productive effort (ootheca); the cockroach cannot resume oogenesis without renewed feeding. The American cockroach still produces oothecae while starved (Kunkel, 1966; Bell

1971)* Bell (1971) monitored protein levels in starved fe­ males of americana. Vitellogenins (proteins used only in reproduction) almost disappeared by the tenth day of starva­ tion, only to increase again when mature oocytes were 33 resorbed. The insect then used this temporary supply of protein* total blood protein levels followed the same pat­ tern. Protein is therefore crucial to oogenesis* P. ameri­ cana sacrifices egg development when starved by sequester­ ing proteins targeted for developing eggs and ultimately by resorbing eggs being formed.

Lipid is also important in oogenesis. Krishnan (1968) found that adult females of B_j_ germanica contained 2 8.9# lipid by weight* oothecae were 3 6 .6# lipid by weight. Most of this lipid concentration is in the form of neutral lipid storage, to be used by the embryo as an energy source. Li­ pid is not considered a critical nutrient in the present study* lipid ingestion was decreased by dilution with dex­ trose, as was protein ingestion (see above), but excess carbohydrate is converted to lipid by germanica (Gordon,

1972). Gordon (1972) also described other nutritional pathways for excess nutrients in cockroaches. Glucose is usually oxidized completely. When present in excess, it is stored as glycogen* when that system is overtaxed, the excess is used to produce triglycerides (see above). If even more glucose is ingested, free glucose accumulates in the blood. Glucose may also be converted into several amino acids, some with the help of bacterial symbiotes (Henry, 1962* Hen­ ry and Block, 1962)* individuals of germanica fed the low protein diet may make up for some of the protein depletion with excess sugars. Amino acids are normally used to con­ struct body tissue and egg protein. Excess essential amino acids are either rapidly oxidized or converted to glucose. At the same time, excesses of both essential and nonessen­ tial amino acids can be converted into ammonia for gluta­ mine, chitin, and purine synthesis. The purine pathway serves as a route for disposal of excess ammonia by conver­ sion into uric acid, which may happen in populations fed the higher protein diets. The cockroach is different from other insects in recycling uric acid with the help of symbiotes

(Brooks and Richards, 1955s Mullins and Cochran, 1975&»bi Valovage and Brooks, 1979)• Early nutritional studies in which the role of sym­ biotes was not clear were made by Melampy and Maynard (1937)* McCay (1938), House (1949a,b), Noland and Baumann (1949, 1951)* Noland, Lilly, and Baumann (1949), and Hilchey (1953)* A comprehensive study was made by Gordon (1959)* who implicated bacterial symbiotes in several pathways. Subsequent work showed symbiotic involvement in amino acid synthesis from glucose (see above), sulfur metabolism (Hen­ ry and Block, i960), vitamin synthesis (Ludwig and Galla­ gher, 1966), and urate metabolism. The use of nitrogenous wastes by the cockroach, with the help of symbiotes, is of particular interest in the present study. Uric acid is not a major excretory product of cockroaches, but is stored in the fat body (Mullins and Cochran, 1972, 1973)* Cochran (1979) has estimated that over 85# of stored urates are found there. High protein in the diet does not increase uric acid concentration in the faeces of americana (ca. 2#)) if uric acid is included in the diet, not all of it is excreted (Mullins and Cochran, 1972). Accumulation of uric acid increases with increased nitrogen intake (Gier, 19^7* Haydack, 1953* Mullins and

Cochran, 1974, 1975a »U; Cochran, 1979* and see below)) in extreme cases, storage includes precipitation into the body cavity (Haydack, 1953)■ Uric acid accounts for up to 12# of dry body weight under normal circumstances (Cochran,

1979)* It is also found in newly-hatched nymphs of amer­ icana, and increases in concentration as the nymphs age

(Cochran, 1979)* The bacterial symbiotes of cockroaches play a signifi­

cant role in urate storage. Brooks and Richards (1955) noted that aposymbiotic (symbiote-lacking) American roaches accumulate more uric acid in the fat body than xenic (sym­

biote-containing) roaches. Urates in xenic German cock­

roaches were 23 # of the dry weight of the fat body; the figure was 48% in aposymbiotic roaches (Valovage and Brooks,

1979)* Urate-containing cells in the fat body (urocytes)' also surround the symbiote-containing cells (mycetocytes)

in P_j_ americana (Bodenstein, 1953) and germanica (Valo­

vage and Brooks, 1979)* The metabolism of uric acid begins with uricase con­ verting uric acid to allantoin. Uricase is also found in S . gregaria. but allantoin is the endpoint of urate metab­ olism in this acridid. Donnellan and Kilby (1967) worked out one symbiote-assisted pathway of urate metabolism in the American roach. Allantoic acid is produced from allan­ toin, and in turn is ultimately converted to glyoxylic acid and ureaj glyoxylic acid is then converted into tartronic semialdehyde, and urea into ammonia and carbon dioxide. Tartronic semialdehyde is converted to glycerate and ulti­ mately into pyruvate, which is injected into the tricar­ boxylic acid cycle. Urates are thus recycled to provide energy from a source that is merely excreted in most other

insects. There is also evidence that uric acid stores in the cockroach may be used in other pathways (Mullins and Coch­ ran, 1975b)- The boll weevil, Antonomus grandis, produces radioactive RNA, DNA, amino acids, and carbon dioxide two 1 ii hours after administration of labelled uric acid-2- C.

Nineteen amino acids were producedj the greatest specificity was shown in valine, threonine, isoleucine, leucine, methio­ nine, tryptophan, cysteine, and serine (Mitlin and Wiygul,

1973)* Uric acid levels decrease just after adult eclosion in both the boll weevil (Mitlin and Wiygul, 1973) and the American cockroach (Mullins and Cochran, 1975a)• which corresponds to the period of great protein need (in 37

S . gregariai Hill, et al., 1968* Mordue (Luntz) and Hill, 1970). Vitellogenin synthesis occurs in the fat body in P. americana (Pan, Bell, and Telfer, 19^9) I "the amino acids used for the synthesis may come from metabolized urates. The effects of different levels of protein intake was first examined by Gier (1947) and Haydack (1953)* Haydack (1953) found optimal diet protein levels of 11-24# in B. germanica and 24-4?# in P^ americana. Protein levels above the optimal range in the American roach caused deposition of white crystals (urates) and ultimately starvation due to lipid or carbohydrate insufficiency. Gier (1947) also blamed the death of P^ americana fed protein-rich diets on lipid insufficiency. Mullins and Cochran (1975a,b) studied the relationship of dietary protein levels to uric acid use in the American cockroach. Newly-eclosed adults were fed diets containing

5, 24, 50 , 79* and 91% protein (Mullins and Cochran, 1975a). Cockroaches fed the 5% protein diet lost 0.13 mg of uric acidj roaches on the other diets gained uric acid, ranging from 0.18 mg on the 24# diet to 1.24 mg on the 79# diet. Weight gain occurred in roaches fed all diets, but most of the increase is explained by increased uric acid storage and not increased body tissue. Some of the weight increase not explained by uric acid may be due to protein-urate com­ plexes in the fat body, to more urocytes produced in the fat body to enclose the additional uric acid, and to extracellular uric acid storage (see Haydack, 1953)* The reaction of P_;_ americana fed the $$ protein diet may be the most interesting for the present study. Total body nitro­ gen and uric acid storage decreased, but consumption and egg production did not, indicating a shift in metabolism to preserve egg production. Mullins and Cochran (19?5b) made further studies on protein depletion. Three diets were fed to newly-eclosed adults. The first diet was cellulose and 5# protein plus salts and vitamins; it was a starvation diet and resulted in high mortality. The other two diets were* dextrin and

5 # protein plus salts and vitamins ("dextrin-plus"), and dextrin only ("dextrin-only"). The percentage of urates in the body decreased in individuals fed both diets, al­ though cockroaches fed the dextrin-plus diet initially in­ creased weight slightly. Seventy percent of the initial weight loss in individuals fed the dextrin-plus diet was due to uric acid loss; after 17 weeks, that figure was 60#.

Cockroaches fed the dextrin-only diet lost 8# of initial weight as urates, and 60# of the weight loss as urates by week 17. Individuals fed both diets were fecund, although adults fed the dextrin-only diet produced more oothecae. A large proportion of total nitrogen loss was due to egg pro­ duction* 52 # in individuals on the dextrin-plus diet, 89# in females fed the dextrin-only diet. Urate mobilization

(metabolism of urates to usable nitrogenous compounds) occurred in individuals fed both diets, but was higher in adults fed the dextrin-only diet. Protein depletion there­ fore results in a mobilization of urate reserves built up during the juvenile instars (see above? Cochran, 1979)i this mobilization is proportional to nitrogen intake and is at the expense of somatic function, since a large pro­ portion of weight loss was due to stored uric acid deple­ tion. CHAPTER V CONCLUSIONS

The results of the present study can now be explained in terms of the level of dietary protein intake and urate storage and mobilization. Individuals of populations fed the low protein diet assimilated less protein than indivi­ duals fed higher protein diets (see above). Weight just af­ ter adult eclosion did not show diet-related differences (Table 11) although less dietary protein resulted in slower development (Table 2). Since weight was not different, it can be assumed that uric acid accumulation and storage from the juvenile stage is also not different. Adult somatic growth also did not show differences associated with diet (see above)s once reproductive activity started, there were diet-related differences. Reproductive investment (Table 16) and output per suc­ cessful ootheca (Table 8) did not show diet-related differ­ ences; as in P_^ americana (Mullins and Cochran, 1975a)» the tradeoff appears to be in less somatic development (urate storage; Mullins and Cochran, 19?5a,b) during reproduction. Individuals of populations fed the high protein diet gained weight after adult eclosion (Table 11); individuals of pop­ ulations fed the medium protein diet gained less weight (Table 11). Weight gain in both populations probably repre­ sents storage of excess ingested protein as uric acid (see above)j the costs of reproduction, in terms of protein re­ quirements, were met by ingested protein. Individuals of populations fed the low protein diet gained weight after a- dult eclosion (Table 11) and produced the first ootheca just as fast as females of populations fed the higher protein diets (Table *+) j a protein deficit is not yet apparent. Prom adult sampling A to adult sampling B, no weight gain occurred (Table 11), and the relative growth rate (Table 12) and both net and gross conversion efficiencies (Tables 13 and 1*0 were negative. Reproductive investment and output kept pace with females fed more protein (Tables 8, 16, 17, 18) due to urate mobilization (see above; Mullins and Coch­ ran, 1975a.b). The protein level of the low protein diet apparently did not provide enough protein for reproduction at sampling B. Urate stores, accumulated during the juve­ nile stage, were probably mobilized to provide the differ­ ence. Other possible sources of protein are the cuticle and muscle tissue, but the urate pathway appears to be able to meet protein requirements, as in americana (Mullins and Cochran, 1975a,b). As a result, urate stores decreased, body weight decreased, some individuals even died, but re­ productive protein needs were ultimately met, and reproduc­ tive investment and output did not suffer in comparison to females of populations fed sufficient protein. Between 42 adult sampling B and C, reproductive investment and output did not show diet-related differences (Tables 8, 16, 17» 18), Females of populations fed the high protein diet had the greatest weight, while individuals of populations fed the medium and low protein diets were progressively lighter (Table 11). Such results can be similarly explained in terms of protein inake and urate mobilization. The 11-24$ "optimal** protein range of Blattella eerman- ica (Haydack, 1953) represents a 50# change in protein con­ tent. Using Haydack's (1953) criteria for the best dietary protein level, mortality and length of developmental period, the three diets used in the present study are not equiva­ lent. The medium protein diet caused the lowest mortality (Tables 1 and 3) and the shortest developmental period (Ta­ ble 2); the high protein diet evoked the next best perfor­ mance, followed by the low protein diet. Distinctions (in­ cluding a peak) within Haydack*s (1953) range do exist. Des­ pite these differences, the similarity of results in view of the 50# difference in protein levels is interesting. Food use did not change in response to protein levels* B_j_ german- ica assimilated less protein and simply functioned with the smaller amount. Developmental rates were lower and mortali­ ty slightly greater, but reproductive performance was not affected* the tradeoff was probably facilitated by urate mobilization. Physiological adaptations buffered Blattella germanica from a huge drop in protein intake, allowing surviving adults to contribute as effectively as females ingesting more protein. BIBLIOGRAPHY

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Wyss-Huber, M., and M. Luscher. 1972. In vitro synthesis and release of proteins by fat body and ovarian tissue of Leucophora maderae during the sexual cycle. J. In­ sect Phvsiol. 18i 689-710. 58 Table 1. Pooled number of survivors at each instar for each diet (juvenile period), and G-test for differences (if significant, diets listed in increasing order by initial! L, low protein} M, medium protein} H, high protein)

Instar Diet G-test Low Medium High

Hatching 1374 1402 1396 G = 3.548 NS 1 1113 1231 1214 G = 9.689** (L,H,M) 2 975 1051 959 G = 7.132* (H.L.M) 3 905 870 884 G = 2.761 NS 4 690 732 744 G = 3.921 NS 5 552 630 622 G = 7.610* (L,M,H) 6 447 567 589 G = 23.734*** (LfM,H) Adults 256 363 364 G = 25.292*** (L,M,H)

Table 2. Days to adulthood, each diet.

Diet Median Range

Low Protein 92.828 73.86?-111.103 Medium Protein 88.797 71.439-93.105 High Protein 85-732 72.303 -98.150

Jonckheere-Terpstra Test for Low^High^ Medium alternative! j* = 1 .749 *, z {< 05) = 1.645. Table 3* Pooled number of female survivors at beginning of adulthood and at first three reproductive efforts for each diet, and G-test for differences a. Reproductive Diet: Effort Low Protein Medium Protein High Protein G-test vs. x (.05.2)^-991

Adult Emergence 16** 23^ 202 G=12.8?l*** I 81 128 105 G=10.905*** all II 52 102 76 G=l6.?19*** (Med>Hi>Low) III 35 6? 5^ G= 10.^6*** b. p Paired comparisons of diets, based on G-test (vs. X ^ ^=3-8*1-1)

Reproductive Effort Low vs. Medium Low vs. High Medium vs. High Adult Emergence G = 12.259*** G = 3.8^5* G = 12.259*** I G = IO.599*** G = 2.797NS G = 10.599*** II G = 15.820*** G = if.118* G = 15.820*** III G = 9.5^2*** G = 3-639NS G = 9-5^2***

NO Table 4. Interval (days) between adult eclosion of females and the appearance of the first ootheca.

Diet Median Range Low Protein 15.028 12.333-18.**29 Medium Protein 13-508 10.588-23.250 High Protein 15-563 9-222-24.130

2 Kruskal-Wallis Test for diet differences (vs. X , nc 5*991) : H = 1.644NS Table 5- Pooled number of offspring produced at first three reproductive efforts for each diet, and G-test for differences a. Diet Reproductive Low Medium High G-test ( v s . 2)=5, Effort

I 1976 3255 2276 G=352.9**# II 1351 2462 1557 G=379.2*** all III 974 1677 1114 G=215.8*** (M>K>L)

Total 5748 9062 6413 G=8 5 8 .6*** (All Efforts) b. Paired comparisons of diets, based on G-test

I G=172.6*** G=19* 9*** G=3l4.4*** II G=204.4*** G=13.7*** G = 327* 3*** III G=113 .5*** 0=8 .8*** G=187.9*** Total G=371-8*** G=l6 .4*** G=252.8*** (All Efforts)

Table 6. Pooled values of number of offspring produced per reproductive effort at first three reproductive efforts for each diet

Reproductive Low Protein Medium Protein High Protein Effort ______

I 25.333 25.233 20.691 II 25.981 24.137 20.221 III 27.829 25.030 21.019 Total 22.992 21.942 18.697 (All Efforts) Page Test for Low>Medium>High alternativei L=56***, 1 (.05.3.'O=5'k 62

Table 7» Pooled values of percentage of successful repro­ ductive efforts at first three reproductive efforts for each diet.

Diet Reproductive Effort Low Protein Medium Protein High Protein

I 83.3 77-5 67.3

II 84.6 76.5 67.5 III 91.4 80.6 73*6

Total 78.4 6 9 .7 6 5 .O (All Efforts) Page Test for Low>Medium>High alternative* L = 42**, 1 (.05,3,3) = k1'

Table 8. Pooled values of number of offspring produced per successful reproductive effort at first three reproductive efforts for each diet.

Diet Reproductive Effort Low Protein Medium Protein High Protein

I 30.400 32.500 30.575 II 30.705 31-564 29.942

III 30.438 31.056 28.564

Total 29.327 31-465 28.758 (All Efforts) Table 9* Medians and ranges (in parentheses) of values of approximate digestibility for each diet at each sampling period, expressed as percentages. a. 1. Juveniles

Instar Low Protein Medium Protein High Protein

1 87-98(82.45-91.33) 85.69(81.43-89.35) 79.37{72.92-87.21) 2 85.35(78.67-88.94) 83-95(78.45-89.95) 77.20(73.03-83.33) 3 86.48(68.71-88.59) 83.88(61.91-87.15) 73-25(47.60-82.93) 4 84.49(78.75-91.6?) 81.86(77*47-84.44) 75.51(72.45-79.76) 5 81.47(77.59-84.21) 80.35(78.50-82.24) 72.49(70.50-82.35) 6 81.03(75.61-83.91) 79-24(75.27-81.19) 71.51(65.39-81.61) 2. Adults

Adult Sampling Low Protein Medium Protein High Protein

A 81.32(76.54-86.00) 76.94(75.53-81.96) 67.94(65.12-94.52) £ 87.69(84.88-88.55) 81.72(65.56-84.91) 72.90(67.78-75.00) C 87.96(85.00-89.29) 83.13(78.57-85.95) 74.45(70.15-76.81) Table 9 (continued). b. Jonckheere-Terpstra Test for Low>Medium>High alternative and Paired Comparisons Tests 1. Juveniles , Paired Comparisons Instar j*a Low vs. Medium Low vs. High Medium vs

1 3.954*** 5.30NS 14.80* 9-50* 2 3.726*** 2.IONS 14.10* 1 2.00* 3 3.802*** 4.00NS 14.00* 1 0.00* 4 4.639*** 5.IONS 17 .10* 1 2.00* 5 3.726*** 3-IONS 14.30* 1 1 .20* 6 3 .536 *** 2.50NS 13.70* 1 1 .20* a. vs. z(.o5)= 1 , 6 5 b. vs. 95% cutoff of 9.226

Adults Adult Sampling J*a Low vs. Medium Low vs. High Medium vs.

A 3 .916*** 6 .00NS 14.40* 8.40NS B 5.285*** 10 .80* 18.90* 6.50NS C 5 .4 7 5 *#* 8.80NS 19.40* 1 0.60* &. vs. Z(.05) = 1,65 b. vs. 95% cutoff of 9*226

•p- Table 9 (continued). c. Page Test for change in Approximate Digestibility with increasing age: 1. Juveniles (decrease) (vs. 1^ g = 777): Low Protein, 854***; Medium Protein, 867***; High Protein, 848***

2. Adults (increase) (vs. 1^ ^ = 128: Low Protein, 134**; Medium Protein,

131**; High Protein, 155***

Os Vn Tattle 10. Medians and ranges (in parentheses) of the Consumption Index for each diet at each sampling period, expressed as mg. consumed per mg. body wt. per day (all weights are dry weights). a. 1. Juveniles

Instar Low Protein Medium Protein High Protein

1 0.420(.289-.776) 0 .388 (.290-.423 ) 0.378(.293-*52^) 2 0.218( .155 -0 0 0) 0.191(.143-.266) 0.204(.102-.279) 3 0.1?5(.129-.283) 0.170(.042-.269) 0 .168{.070-.298) 4 0.103(.062-.l4l) 0.090(.071-.155) 0.147(.083-.196) 5 0.100(.057-.184) 0.131(.071-.186) 0.109(.074-.155) 6 0.086(.059-.124) 0.09^(.O83 -.I3 6 ) 0.095(•078 -,126)

2. Adults

Adult Sampling Low Protein Medium Protein High Protein

A 0.073(.O63 -.IOO) 0.102(.083-*133) 0.098(.024-.383 ) B 0.095(.084-.131) 0 .093(.059-*109) 0.095(.050-.104) C 0.082(.067-.1^0) 0 .084(.084-.121) 0 .069(.044-.075) b. Kruskal-Wallis and Paired Comparisons Tests for diet differences

1. Juveniles - Only (Medium

2. Adults - Sampling A: (Low

CN Os Table 10.(continued). c. Page Test for changes with age

1. Juveniles (decrease) (vs. 1^ = 777)} Low Protein, 893***; Medium Protein, High Protein,*875***

2. Adults (vs. 1^ j = 128: Low Protein (increase), 132*; Medium Protein (decrease), 133*; High Protein (decrease), 128* Table 11. Medians and ranges (in parentheses) of values of cumulative production (in mg. dry weight) for each diet at each sampling period (ten individuals sampled per diet) a. 1. Juveniles

Instar Low Protein Medium Protein High Protein

1 0.449( .206-.952) o.5 l5 ( .1^-2-.652 ) 0.504(.204-.594) 2 0.795(-498-1.186) 0.837(.702-1.060) 0.789(.354-1.296) 3 1 .579(.812-2 .016) 1.447(. 776-2.042) 1*585(-898-2.172) 4 4.147(2.612-5-280) 4.756(1.834-8.278) 3.018(1.574-5.310) 5 7.350(5.128-12.268) 8 .024 (5 *386 -1 4 .575 ) 7.842(5.770-10.880) 6 1 3 0 2 9(10.086-18.712) 15-929(8.426-18.972) l6 .96K l l . 738 -i9.494 ) 2. Adults

Adult Sampling Low Protein Medium Protein High Protein

A 16.023(18.560-29.503) 16.530(9.720-25.226) 16.025(10.928-31.204) B 16.063(12.814-19.334) 19.772(16.660-23.345) 23.492{18.597-27.210) C 20.450(13.425-28.834) 23.727(19.574-28.450) 29.610(20.209-39.292)

CN 00 Table 11 (continued). b. Juveniles: Kruskal-Wallis Test for diet differences: none at any age (P<.05)

Adults: Jonckheere Terpstra Test for High>Medium>Low differences and paired comparisons:

Adult Paired Comparisons Sampling J*e Low vs. Medium Low vs. High Medium vs. High

A 0 .266ns B 4.48?*** 9.50* 1 6 .60* ?.IONS C 3.422*** 3.20NS 1 3 .00* 9.80* a. vs. Z^ = 1.65 b. vs. 95% cutoff of 9.226 c. Page Test for increased cumulative production with increased age in adults (vs. 1( q

Table 12. Pooled values of relative growth rate (in mg. gained per mg. wt. (dry wts.)),each sampling period, for each diet

1. Juveniles Instar Low Protein Medium Protein High Protein 2 .051 To52 .056 3 .0 45 .03 *+ .0*+7 4 .022 .032 .025 5 .026 .026 .030 6 .025 .029 .034

2. Adults Adult Sampling Low Protein Medium Protein High Protein A .010 Took .002 B -.007 .004 .007 C .003 .002 .003

Juveniles - Page Test for High Medium Low alternative (vSl 1 (.05.3,5) = 66)1 L = 67'5*

- Page Test for decreased relative growth rate with increasing age (vs. l (.05,5,3) = 150T. L = 157** 71

Table 13. Pooled values of conversion efficiency of ingested food (as percentages, based on dry weights) for each diet

1. Juveniles Instar Low Protein Medium Protein High Protein 2 23.5 25.1 2B .1 3 25.1 20.if 28.8 if 20.8 33-0 18.0 5 23.6 20.3 26.5 6 27.2 29.3 35.**

2. Adults Adult Sampling Low Protein Medium Protein High Protein A 13.2 378 178 B - 7.0 if.if 7 .7 C 3* ^ 2.2 if.5

Diet differences: _ - Juveniles: Friedman Test (vs. X { o\=5*991) 1 S = 2.800NS. 2 - Adults: Friedman Test (vs. X , o,i-5*991) * S = 0.667 NS. I * wj,^ j 72

Table 14. Pooled values of conversion efficiency of (digested) food into body substance (as percentages, based on dry weights) for each diet

1. Juveniles Instar Low Protein Medium Protein High Protein 2 27.7 31.1 357! 3 29.8 24.5 &0.9 4 24.7 40.5 23.8 5 29.2 25.2 3 6 .I 6 33-8 37*2 49.3

2. Adults Adult Sampling Low Protein Medium Protein High Protein A --- 1773 ----- 479------2 . 4 ------B - 8 .0 5 .5 10.6 C 3*9 2.7 6.1

Diet differences! p “ Juveniles; Friedman Test (vs. X , nc ~ 5*991) 1 S = 2 . 800NS. 2 - Adults: Friedman Test (vs. X / - 5.991) t S ^ “07667NS. (.05.2) Table 15. Medians and ranges (in parentheses) of oxygen consumption (expressed as i/1 O^/mg. dry wt./day for each diet at each sampling (one individual sampled per diet) a. 1. Juveniles

Instar Low Protein Medium Protein High Protein

1 135.20(53.52-^63.52) 113.01(44.65-298.85) 128.38(17.51-592.04) 2 88.99(^8.68-32?,93) 80.32(31.72-237.82) 87.59(54.91-173.94) 3 56.78(36.62-115.01) 64.02(29.58-212.84) 62.70(37.27-252.40) 49.62(25.81-111.17) 55.06(35.88-171.48) 54.91(28.67-146.16) 5 47.63(33.14-68.01) 47.55(26.03-81.09) 39.22(18.00-72.63) 6 39.46(32.00-59.86) 44.03(13.60-88.82) 35-06(19.18-66.34) 2. Adults

Adult Sampling Low Protein Medium Protein High Protein

A 48.12(35*72-81.53) 55.30(32.93-111.08) 40.20(28.03-83*04) B 46.17(25.98-84.12) 36.07(18.56-56.83) 23.5507.79-52.^9) C 45.12(21.09-65.71) 41.82(23.82-86.72) 21.65(7.23-37.98) Table 15 (continued). b. Diet differences:

- Juveniles: Kruskal-Wallis Test (vs. X2^ = 5*991) all < 2.681

- Adults: Jonckheere-Terpstra Test (vs. Z^ = I.6 5 )

Adult Sampling J* _____Order______

A 2.091** Mediura>Low>High B 3 .^98*** Low>Medium>High C 2.852*** (Medium = Low)>High

Page Test for Medium>Low>High alternative (vs. 1^ 3 ,3 ) = ^'1)* L “ ^1* c. Page Test for decrease in respiratory rate with increased age

- Juveniles (vs. 1/ £ = 77?)* Low Protein, 869***} Medium Protein, 828***t (.05 ,of10j High Protein, 855 *** - Adults (vs. 1, l M = 128): Low Protein, 12^NS; Medium Protein, 127NS; (•0D 1 J*LU ) High Protein, 130* 75

Table 16. Pooled values of reproductive investment (expressed as total mg. dry weight of oothecae produced per mg. dry weight of insect per day)for each diet

Adult Sampling Low Protein Medium Protein High Protein A .018 .021 .027 B .028 .015 .019 C .016 .012 .012

2 - Diet differences: Friedman Test (vs. X / n< = 5-991) (S'for tied values): S '= 1.151NS. * ‘ J } - Page Test to test for decrease in reproductive effort with increasing age (vs. 1 / ~ = 4l) : L - *H*. \ Jt j ) 76 Table 17- Reproductive biomass produced per day as a percentage of total food ingested per day (dry wts.), each sampling period, for each diet. Adult Sampling Low Protein Medium Protein High Protein A ” 2 4 7 3 3 ----- 20.59 27.55 B 29.^7 16.13 20.00 C 19.51 14.29 17.39

Freidman Test for Diet differences (vs. X = 5.99D 1 S - 4.667NS. (.05,2) Page Test for a decrease with increasing age (1 41) ( L - 41*. (.05,3,3)

Table 18, Reproductive biomass produced per day as a percentage of total food digested per day (dry wts.), each sampling period, for each diet.

Adult Sampling Low Protein Medium Protein High Protein A 30.51 23792 40.30 B 33-73 19.74 27.54 C 22.22 17-14 23-53 Friedman Test for diet differences (vs. (.0 5 ,2 ) = 5.991) . S = 4.66?NS. Page Test for a decrease with increasing age (1/ ^ -v 41) : L = 41*. (.05,3,3J Table 19* Medians and ranges (in parentheses) of dry weights of second-effort oothecae (in mg.) for each diet.

Low Protein Medium Protein High Protein 11.309(7-358-13-980) 9.3^1(5-928-13.268) 11.088(7.572-22.808)

2 Diet differences 1 Kruskal-Wallis Test (vs. X / ne = 5.991) (H* for tied values) t H* = 3-259N3: ’3t }

Table 20. Pooled values of number of oothecae per female per day for each population sampling period, each diet. Adult Sampling Low Protein Medium Protein High Protein A .032 .039 .040 B .040 .037 .037 c .031 .030 .030

2 Diet differences! Friedman Test (vs. X , 0 x = 5*991) (S1 for tied values) : S = 0.716NS. K } Page Test for decrease in number of efforts with increasing age (vs. 1 ^ , 05 ,3 ,3 ) = ^1 ) ' L = Table 21. Food use values of selected insect species (other indicesi Cl, consumption index? R, respiration t ingestion; RA, respiration * assimilation).

Net Gross Approx. Conv. Conv. Other Order. Species Food Dig.(%) Eff.(%) Eff.(%) Indices Source Lepidoptera 1 .Bertha armyworm Cruciferae 50 4 5 21 Bailey (1976) (Mamestra configurata) Rape (Brassica 40.7 52.7 21.3 — — — Bailey & napus) Singh (1977) 2.Southern armyworm Cranberry 48.5 « » h 33.5 » ■ — Crowell (1941) (Prodenia eridanea) 18 plants 36.3- 1 6 .2- 8.4------Soo Hoo & 72.6 56.8 38.1 Fraenkel (1966) 3.Phalera bucephala Hornbean 35.0 Cl Evans (1939) 0.233 4.Gypsy moth Artificial 49.7 36.9 17.5 Barbosa & (Lymantria dispar) Greenblatt (1979) 5.Malacosoma neustria Wi Ho w 34 Cl Evans (1939) 0.326 6 .Aglais urticae Nettle 25-7 ... B B Cl Evans (1939) 0.581 — — _ ^ _ 7.Cabbage butterfly Cabbage 36.3^ CI a Evans (1939) (Pieris brassicae) 39.1 0.233 0.188 R=2l% 8 .Cyclophragma leucosticta ? 38c 17° Mackey (19?6) 9.Monarch butterfly Milkweeds 46-50 — ------Mathavan & (Danaus chrysippus) Bhaskaran (1975) Milkweed 45 4l Mathavan & Bhaskaran (1975) Table 21 (continued).

Net Gross Approx Conv. Conv. Other Order, Species Food Dig.(# Sff. {%) Eff.[%) Indices Source

D. chrysippus, continued Milkweed 46 c 46.6c 24.6c Schroeder 4ld 20.5 d (1976) 10.Miner •? 72.6 20.6 15.0 — New (1979) (Labdia spp.) 1 1 .Range caterpillar Artificial 54 52 23 Cl Schowalter, (Hemileuca oliviae) •3 3 -‘^0 Whitford, & Turner (1977) 12.Cecropia moth Box elder 36 . 6c 53-lc 19.4c -— Schroeder (Hyalophora cecropia) 29.2d 5 6 .2d 1 6 .4d (1971) 13-Cecropia moth Lilac 34c 48.0c 16,2c -- Schroeder (Platysamia cecropia) 31 d 52d 19d (1972) l4.Pachvsohinx modesta Aspen 33-3c 57c 19c -- Schroeder 29.7d 46 d 13.7d (1973) 15‘Milkweed tiger moth Milkweed 34.3 42.6 14.6 Schroeder (Euchaetias egle) (1976)

Orthoptera 16.House cricket Synthetic 62 15.5- 25-27 — Woodring, Roe, (Achaeta domesticus) 16.5 & Clifford (1977) ? 64.7 — — — — — ——— Crossley & Van Hook (1970) 1?.Two-striped grasshopper Alfalfa, 41-47 35-44 16,5 Bailey & (Melanoplus bivittatus) corn, let- Mukerji tuce, radish (1976) 18.Red-legged grasshopper Alfalfa, 37-48 33-44 17 — Bailey * (Melanoplus femurrubrum) corn, let­ Mukerji tuce, radish (1976) no Table 21 (continued).

Net Gross Approx. Conv. Conv. Other Order, Species Food Dig.{%) Eff.{%) Eff.(%) Indices Source

19•Migratory grasshopper Wheat, oats, 32 2? wes.wh. -- -- Smith (1959) (Melanoplus bilituratus) western gr., wheat grass 32 wheat, 38 oats 20.Grasshopper Creosote- 22c 4lc 9c R,13 Mispagel (Bootettix punctatus) bush RA.59 (1978) 21.Dusky grasshopper Grasses 26.1c 40.6c 10.6c -- Bailey & (Enloptolophus sordidus Riegert costalis) (1972) 22.Rock grasshopper Moss & 20.1c 30.0c 6.0c RA,?0 Duke & Cross- (Trimerotropis saxatilis ) lichen ley (1975)

Coleoptera 2 3 .Black carpet beetle Artificial 70.2 32.6 22.9 Cl Baker (1975) (Attageuus megatoma) 0.115 24.Mealworm Bran 46.3 -- -- Evans & (Tenebrio molitor) Goodcliffe (1939) 25*Boll weevil 49c -- -- McGovern, et (Anthonomus grandis) 66. Od al. (1976) 26.Yellow poplar weevil Lirioden- 51c 22c 11c RA.78 Van Hook & (Odontopus calceatus) dre.n tuli- 54d 19d lOd Cl, Dodson pitera 0.970 (1974)

Homoptera 27*Spittlebug 41.6 36.9 15.4 Hinton (1971) (Neophilaeuus lineatus) Table 21 (continued).

Net Gross Approx. Conv. Conv. Other Order. Species Food Dig.(*) Eff.(%) Eff.{%) Indices Source 28.Pea aphid Sap of 83.8 56.6 4?. 4 R,36.6 Randolph, et (Acyrthosiphon pisum) Alaska var. RI.43.6 al, (1975) peas

29.5talia .major (predator) 51 -- Fewkes (i960) 30.Rhodnius prolixus human 22.1 -- Friend, Chay, blood & Cartwright (1965) Hvmenoptera 31.Sawfly Scots 14 Lars son & Ten- (Neodiprion sertifer) pine ow (1979) 32.Sawflies pines 17 33 5 Cl Fogal <197*0 (4spp.f Neodiprion. (4 spp.) 1.9 Diprionl

a. fed young leaves b. fed old leaves c. energy budget study d. biomass budget study Table 22. Averages and standard deviations of food use values of selected insect orders from Table 21 and Blattella germanica.

a. Order Consumption Approximate Net Conversion Gross Index Digestibility Efficiency Conversion Efficiency

(mg/mg/day) — m ------— w

Lepidoptera 0 .34 3±.148,n=5 42.29111.47,n=19 43.54+ 9.67,n=12 19-67+5.21,n=13 Orthoptera 38.66+16.I8,n=9 34.51+8.31,n=10 l4.30+2.57,n=7 Coleoptera 0,5^31■605,n=2 59*13110.96, n=4 25-80+9-62,n=2 l6.45+9.12,n=2 Homoptera 62.70+29.84,n=2 46.75±13-93»n=2 31 .40 +2 2.63 ,n=2 Hymenoptera 1.9,n=l 15.50+1.46,n=2 33,n=l 13-5514.52,n=2 Hemiptera 36.55±4.52,n=2

Total 0.588 +.771 ,n=8 42.90+4.03,n=36 38.73+3-25,n=2? 19.62+3.24,n=27

b. B. germanica Consumption Approximate Net Conversion Gross study Index Digestibility Efficiency Conversion Efficiency

McCay (1938) 31.4 Brooks (1965) -- 51 C02,64 cont. Woodland, et al.(1968) 83 54(a) 45(a) Present Study 0.150+.098,n=27 80.1915*54,n=27 32.66+7 .38(b),n=15 25.74+4.85(b), n-15 a. 3~5th instar males b. juveniles only

00 to Table 23. Compensations in food use (+, positive evidence for mechanism; negative evidence for mechanism).

Order, Species Variable Approx. Dig.- Approx. Dig.- Source Consump.Index Net Conv. Eff.

Lepidoptera 1.Bertha armyworm Diet Bailey (1976) (Manestra cenfigurata) 2.Southern armyworm Diet + Soo Hoo & (Prodema endania) Fraenkel (1966) 3.Gypsy moth Diet + + Barbosa & (Lymantria dispar) Greenblatt (1979) 4.Sugarcane borer Parasitiza- + Brewer & (Diatraea saccharalis) tion King (1978) 5-Clothes moth Relative + - Chauvin & (Tinea pellionella) Humidity Gueguen (1978) 6.Tobacco hornworm Insecticide + - Dahlman (Manduca sexta) (1977) 7.Celerio euphorbiae Dilution + House (1965) 8.Pale western cutworm Diet + Kasting & Mc­ (Agrotjs orthogonia) Ginnis (1959) 9.Black cutworm Insecticide + Reese & Beck (Agrotis ipsilon) (1976a) Insecticide Reese & Beck (1976b) 10.Soybean looper Diet +? Kogan & Cope (Pseudoplusia inchideus) (197*0 11.Armyworm Rations Mukerji & (Pseudaletia unipuncta) Guppy (1970)^ Table 23 (continued).

12.Monarch butterfly Diet Mathavan & (Danaus Chrvsippus) Bhaskaran (1975) Rations Mathavan & Mathukrishnan (1976) 13‘Cherry scallop moth Ration Schroeder (Calocalpe undula) (1975) 14,Cecropia moth Food Scriber (Hyalophora cecropia) Moisture (1977) 15.Cabbage butterfly Nitrogen Slansky & (Pieris rapae) Content Feeny (1977) 16.Cabbage butterfly Insecticide Turunen (Pieris biassicae) (1977) Orthoptera 17.Two-striped grasshopper Diet Bailey & (Melanoplus bivittatus} Mukerji (1976) Dilution McGinnis & Kasting (1966) 18.Red-legged grasshopper Diet Bailey & (Melanoplus femurrabrum) Mukerji (1976) 19>Lesser migratory g.h. Dilution McGinnis & (Melanoplus sanguinioes) Kasting (1967) 20.Migratory grasshopper Diet Smith (1959) (Melanoplus bilituratus) 21.Migratory locust Dilution Dadd (I960) (Locusta migratoria) Diet Hoekstra & Beenakkers (1976) Table 23 (continued).

22.Desert locust Dilution + + Dadd (I960) (Schistocera gregaria) 23.Poecilocerus pictus Rations + ? Muthukrishnan & Delvi (197*0

Coleoptera 24-.Black carpet beetle Diet + Baker (1975) (Attagenus megatoma) 25'Carpet beetle Diet +? Baker & (Anthrenus flavipes) Schwalbe (1975) 26.Colorado potato beetle Diet + Latheef & (Leptinotarsa decemlmeata) Harcourt (1972) 27.Flour beetle Pupal wt.at + Medrano & (Tribolium casaneum) 21 days Gall (1976) 28.Flour beetle Diet + Waldbauer & (Tribolium confusum) Bhattacharya (1973)

Hymenoptera 29'Sawflies, ^ spp. Diet 2+ Fogal (1974) (Neodjprion. Diprion) 2-

\0 r 0 . Table 24. Age or size-related trends in food use indices (+, increase; decrease; 0, unclear).

Gross Consump. Approx. Net Conv. Conv. Oxygen Order. Species Index Dig. Eff. Eff. Consump. Source Leipdoptera 1.Bertha armyworm + Bailey (1976) fManestra configurata) 0 + + Bailey & Singh (1977) 2.Pale western cutworm 0 Kasting & Mc­ (Agrotis orthogonia) Ginnis (1959) 3.Armyworm - Mukerji & (Fseudaletia unipuncta) Guppy (1970) 4.Soybean looper 0 - Kogan & Cope (Pseudoplusia includens) (1974) 5.Tobacco hornworm Dahlman (Manduca sexta) (1977) 6.Gypsy moth Edel’man fLvmantria dispar) (1963) 7.Phalera bucephala Evans (1939) 8.Cvclophragma leucostica 0 Mackey (1976) 9.Monarch butterfly + + - Mathavan & (Danaus chrvsippus) Bhaskaran (1975) 10.Range caterpillar 0 0 Schowalter, (Hemileuca oliviae) Whitford, & Turner (1977) 11.Silkworm 0 - Hiratsuka (Bombyx mori) (1920; in Waldbauer, 1968) Table 24 (continued).

Gross Consump. Approx. Net Conv. Conv. Oxygen Order. Species Index Dig. Eff. Eff. Consump. Source Orthoptera 12.Two-striped grasshopper + 0 Bailey & (Melanoplus bivittatus) Mukerji (1976) 13-Red-legged grasshopper + 0 Bailey & (Melanoplus femurrubrum) Mukerji (1976) 14.Migratory grasshopper + Smith (1959) (Melanoplus bilituratus) + Smith (1959) 15- + Smith (1959) 15-Dusky grasshopper Bailey & (Encoptolophus sordidus Riegert costalis) (1972) 16.Migratory locust Beenakkers & (Locusta migratoria) Broek (1974) Beenakkers, Meisen, & Scheres (1971) Clarke (1957) Hoekstra & Beenakkers (1976) 17.Desert locust Davey (1954) (Schistocerca gregaria) 18.Paddy field grasshopper 0 0 Delvi & Pan- (-Qxya velox) dian (1971) 19-Poecilocerus pictus Delvi & Pan- dian (1972) 20.Rock grasshopper Duke & Cross-Qo (Trimerotropis saxatilis) ley (1975) ^ Table 24 (continued)

Gross Consump. Approx. Net Conv. Conv. Oxygen Order, Species Index Dig. Eff. Eff. Consump. Source

21.Boottetix punctatus Mispagel (1978) 22.0rchelimum fidicinum 0 Smalley (i960)

Coleoptera 23.Melasoma collaris 0 Hagvar (1975) 24.Colorado potato beetle + Latheef & (Leptinotarsa decemlineata) Hareourt (19?2) 2 5 .Flour beetle - 0 0 0 - Medrano & (Tribolium casaneum) Gall (1976) 26.Rice weevil 0 + - Singh, Camp­ (Sitophilus oryzae) bell, Sinha (1976)

Hemiptera 27 .Stalia ma.ior + Fewkes (i960) 28.Rhodnius prolixus 0 Friend, Chay, & Cartwright (1965) 29-Bed bug Johnson (i960) (Cimex lectularius)

Diptera 30.Blow fly Hanski (1976) (Lucilia illustris)

00 00 Table 24 (continued)

Gross Consump. Approx. Net Conv. Conv. Oxygen Order, Species Index Dig- Eff. Eff. Consump. Source Homoptera 31.Spittlebug Hinton (1971) (Neophilaenus lineatus) 32 .Pea aphid 0 Randolph, (Acyrthosiphon pisum) Randolph, & Barlow (1976) Acarinae 33.Mite 0 + Stepien & (Caloglyphus berlesei) Rodriguez (1972)

CD '•O 90 250

2 2 5 .

2 0 0 .

CO § 125 . > 3 100_ CO

25

T® O c III H © 3 © Reproductive Effort £ Ui

Figure 1. Pooled number of adult survivors in popula­ tions fed each diet through the first three repro­ ductive efforts. (Solid line, high protein diet; dashed line, medium protein diet; dotted line, low protein diet.) Figure 2. Pooled number of offspring from populations from offspring of number Pooled 2. Figure Offspring medium protein diet; dotted line, low protein diet.) protein low line, dotted diet; protein medium fot. Sld ie hg poende; ahd line, dashed diet; protein high line, (Solid efforts. fed each diet through the first three reproductive three first the through diet each fed 0 0 9 3 0 5 5 3 0 5 4 1 0 0 8 1 0 0 5 2 0 0 2 3 0 5 8 2 2150 0 5 7 . . _ _ . . . . _ I erdcie Effort Reproductive III

91

92 100_

9 5_

9 0.

8 5 .

75 .

■o 70-

6 5 .

5 5 „

II III Reproductive Effort

Figure 3* Percentage of reproductive efforts, pooled for populations fed each diet, that yield offspring, through the first three reproductive efforts. (Solid line, high protein diet; dashed line, medium protein diet; dotted line, low protein diet.) Approximate Digestibility l%l 0 6 0 8 5 6 70 90 5 8 75 ...... Figure reas, medium proteindiet; solid white areas, low proteindiet). each (Cross-hatcheddiet. areas, high proteindiet; dotted a- (expressedas apercent)each atsampling for populations fed k. Medians ofMedianspooledvalues of approximate digestibility • * Instar nraig Age Increasing 6 J L / \ dl Sampling Adult

B ------«t ♦ • • • • • • • • • • • • vo % Consumption 100 50. Figure5* Composite theof fate of ingested (i.e.,food food use) m duringthe lifetimeindividualsof fedthe high protein diet. Respiration mtc Prod. omatic nraig Age Increasing er. Prod Reoro. < a a O « w X E (0 O)

o o c (0 3 E a c o % Consumption Figure 6.Figure Composite of thefate ofingested (i.e.,food food use) duringthe lifetimeindividuals of fed the medium protein diet. oai Pro Somatic Respiration nraig g > Age Increasing

Approximate Digestibility

% Consumption Figure7* Composite theoffate of ingested food (i.e., food use) duringthe lifetimeindividualsof fedthe low protein diet. Respiration oai Prod. Somatic Increasing A ge ge A Increasing >

Consumption vo Ov