Experimental Ecology of Dryas Octopetala Ecotypes. V. Field Photosynthesis of Reciprocal Transplants Author(S): J
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Nordic Society Oikos Experimental Ecology of Dryas octopetala Ecotypes. V. Field Photosynthesis of Reciprocal Transplants Author(s): J. B. McGraw Reviewed work(s): Source: Holarctic Ecology, Vol. 10, No. 4 (Nov., 1987), pp. 308-311 Published by: Blackwell Publishing on behalf of Nordic Society Oikos Stable URL: http://www.jstor.org/stable/3682566 . Accessed: 16/11/2011 18:14 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Blackwell Publishing and Nordic Society Oikos are collaborating with JSTOR to digitize, preserve and extend access to Holarctic Ecology. http://www.jstor.org HOLARCTICECOLOGY 10: 308-311. Copenhagen1987 Experimentalecology of Dryas octopetalaecotypes. V. Field photosynthesisof reciprocaltransplants J. B. McGraw McGraw,J. B. 1987. Experimentalecology of Dryas octopetalaecotypes. V. Field photosynthesisof reciprocaltransplants. - Holarct.Ecol. 10: 308-311. The response of photosynthesisto reciprocaltransplanting was measuredin 1980, 1981, and 1986in two genetically-distinctpopulations of Dryasoctopetala, a circum- polar dwarf shrub. Contraryto expectation, photosyntheticrates were lowest in "home"environments and highestin "foreign"sites. Also, "native"plants had lower photosyntheticrates than "alien"plants. A "rapid-transplant"experiment showed that the observedpattern was not causedby environmentaldifferences between sites at the time of measurement,but ratherby the long-termresponse of transplantsto the environment.These results,in combinationwith the resultsof an analysisof the fitnessresponse to transplanting(published elsewhere), caution against assuming that a positive relationshipalways exists between photosyntheticrate and fitness. Inte- gratedmeasures of carbongain may be more appropriateas measuresof plant per- formance. J. B. McGraw, Dept of Biology, West VirginiaUniversity, P.O. Box 6057, Mor- gantown, WV26506 6057, USA. ation and failed to find it 1. Introduction (but see McNaughton et al. 1974, Kemp et al. 1977, and Clough et al. 1979). Of the physiological traits that vary among plant pop- The majority of studies demonstrating population- ulations, primary ecological importance is often attri- level variation in photosynthesis have been carried out buted to photosynthesis (Hiesey and Milner 1965). This in "common environments" (usually greenhouse or assumption seems justified by the central role of photo- growth chambers). The more powerful reciprocal trans- synthesis in the carbon economy of plants (Mooney plant design (Clausen et al. 1940) allows determination 1972). If plant growth rate is defined as net dry weight of components of phenotypic variance among popula- gain per unit time, net photosynthesis (on a whole-plant tions. Mooney and Billings (1961) simulated such a re- basis) for the same time period is proportional to plant ciprocal transplant design in a growth chamber study growth rate (Osmond et al. 1980). Moreover, growth with arctic and alpine populations of Oxyria digyna. Al- rate is often positively correlated with fitness, at least in though a formal analysis of variance was not presented, theory (McGraw and Wulff 1983). both genetic and environmental effects on net pho- Numerous studies have demonstrated the existence of tosynthetic rate were evident. A simulated reciprocal variability among populations in photosynthetic prop- transplant was also carried out in growth chambers with erties. Populations vary with respect to effects of tem- bog and alpine populations of Ledum groenlandicum perature (Fryer and Ledig 1972, Machler and Nos- (Riebesell 1981), with similar results. However, pho- berger 1977, Teramura and Strain 1979), light (Holm- tosynthetic responses of plants grown under controlled gren 1968, Gauhl 1976, Teramura and Strain 1979), and conditions do not necessarily correspond to their re- drought (Al-Ani et al. 1972) on leaf photosynthetic sponses in field-grown plants, as has been clearly dem- rates. Fewer published studies have sought such vari- onstrated with C3 and C4 tidal marsh species (DeJong et Accepted 25 May 1987 ( HOLARCTIC ECOLOGY 308 HOLARCTIC ECOLOGY 10:4 (1987) al. 1982). It is therefore surprising that photosynthetic snowbed sites (n= 10 for each treatment). Transplants to measurements have not been reported for field recipro- "home" sites for each ecotype served as controls for cal transplants, where native and alien plants are ex- transplants to alien sites. Some soil was transferred with posed to normal seasonal changes in the environment each plant to minimize root damage. A zone 2-4 cm and multifactorial differences between sites. wide was cleared of vegetation around each trans- In the present study, I determined the photosynthetic planted individual. The surrounding plant community response of ecotypes to a field reciprocal transplant ex- was otherwise undisturbed. Plants were watered once periment with the arctic shrub, Dryas octopetala L. (Ro- immediately following transplanting and then allowed saceae). A concurrent study showed that survival, to become established. growth and potential seed production (i.e., collectively over eight growing seasons was approximating fitness) 2.3. Photosynthesismeasurements highest for ecotypes transplanted to home sites (McGraw 1982, McGraw and Antonovics 1983a, 1983b, Maximum photosynthetic rate was measured on leaves McGraw, in press). My null hypothesis was that maxi- of D. octopetala with a portable apparatus designed to mum photosynthetic rates at peak season would parallel deliver 14CO2 to an intact leaf enclosed in a cuvette the observed fitness response to transplanting, i.e., (Shimshi 1969, Tieszen et al. 1974). Photosynthetic maximum photosynthetic rates would be highest for measurements were made by enclosing a leaf in the cu- plants grown in home sites. Maximum photosynthetic vette and exposing the leaf to labelled air for 30 s. Leaf rate was chosen as an index of the photosynthetic re- discs of known area were then excised from the labelled sponse since it sets the theoretical limit of photosyn- leaf and immediately placed in a mixture of 0.5 ml thetic yield (Avery 1977). methanol and 0.5 ml phenethylamine in scintillation vials (Bigger and Oechel 1982). The methanol-pheneth- ylamine mixture killed the tissue, preventing release of assimilated 14C through respiration, and dissolved 14C- 2. Methods labelled products of photosynthesis. The samples were stored for counting at a later date. 2.1. Studysite and plant materials Temperature and light measurements were made in Reciprocal transplants and photosynthetic measure- order to ensure that the field exposure technique would ments were conducted near Eagle Summit at mile 106 give consistent estimates of maximum photosynthetic on the Steese Highway in an unnamed mountain range 14CO2 uptake. All photosynthesis measurements were of interior Alaska (65o25'N 145'30'W; elevation made on leaves oriented perpendicular to direct incom- 1050 m). The 6 ha study site encompassed a snow-free ing irradiance. No measurements were made on leaves ridgetop (fellfield) and an adjacent area of late-lying exposed to a photosynthetic photon flux density of less snow (snowbed) (see McGraw and Antonovics 1983a or than 1000 itmol m-2 s-1. Air temperatures varied less Miller 1982, for detailed description). Dryas octopetala, than 4'C over the course of any one experiment. Meas- a circumpolar, dwarf shrub species, is found throughout urements of the time course of leaf temperatures for a the snowbank study site. However, two distinct eco- leaf enclosed in the leaf cuvette showed that leaf tem- types reach greatest abundance in snowbed and fellfield perature increased only 1.45'C after 30 s. Therefore zones, with very little overlap in distribution (McGraw temperature effects on photosynthetic rates during the and Antonovics 1983a). In a narrow strip between the measurement interval were presumed to be negligible. fellfield and snowbed, intermediate forms may be found Samples were prepared for counting by the procedure which are genetically different from either extreme of Bigger and Oechel (1982) and counted with a Beck- form (McGraw and Antonovics 1983a). Fellfield plants man LS-250 Liquid Scintillation Counter. Due to have small (5-15 mm) deciduous leaves and a compact quench problems, every sample was spiked with a mat-forming growth habit. Snowbed plants have larger known radioactivity to determine a counting efficiency leaves (15-50 mm) that often persist and function pho- for each sample. Photosynthetic rates were then calcu- tosynthetically for two growing seasons. The growth lated according to Tieszen et al. (1974). habit of snowbed plants tends to be more clonal than Maximum photosynthetic rate was measured on Au- that of the fellfield ecotype, with lower shoot densities gust 10, 1980 and August 3, 1981 on adult D. octopetala and extensive adventitious rooting from the woody plants in each of the populations which were trans- stems. The snowbed ecotype is found primarily in Al- planted reciprocally in 1979. The 1980 measures were aska and probably evolved from the fellfield form dur- performed on 5 replicate leaves