Field Measurements of Photosynthesis and Leaf Growth Rates of Three Alpine Plant Species
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Utah State University DigitalCommons@USU All Graduate Theses and Dissertations Graduate Studies 5-1973 Field Measurements of Photosynthesis and Leaf Growth Rates of Three Alpine Plant Species Douglas A. Johnson Utah State University Follow this and additional works at: https://digitalcommons.usu.edu/etd Part of the Ecology and Evolutionary Biology Commons, Environmental Sciences Commons, and the Plant Sciences Commons Recommended Citation Johnson, Douglas A., "Field Measurements of Photosynthesis and Leaf Growth Rates of Three Alpine Plant Species" (1973). All Graduate Theses and Dissertations. 6269. https://digitalcommons.usu.edu/etd/6269 This Thesis is brought to you for free and open access by the Graduate Studies at DigitalCommons@USU. It has been accepted for inclusion in All Graduate Theses and Dissertations by an authorized administrator of DigitalCommons@USU. For more information, please contact [email protected]. FIELD MEASUREMENTS OF PHOTOSYNTHESIS AND LEAF GROWTH RATES OF THREE ALPINE PLANT SPECIES by Douglas A. Johnson A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE 1n Range Science (Plant Ecology) Approved: UTAH STATE UNIVERSITY Logan, Utah 1973 ii ACKNOWLEDGEMENTS Grateful acknowledgement is made to Dr. Martyn M. Caldwell for his generous support in terms of time, ideas, and the prepara tion of this thesis. Appreciation is also extended to Drs. Herman Wiebe and Donald Sisson for their assistance in providing valuable suggestions pertaining to the fieldwork for this thesis. The assis tance of Kathryn Johnson, Russel 1 Moore, James Ehleringer, Dennis Ballinger, Rita Belserene, as well as the station support of the Institute of Arctic and Alpine Research, University of Colorado is sincerely appreciated. This study was supported by the U. S. National Science Foundation as part of the U. S. Tundra Biome of the International Biological Program. Douglas A. Johnson iii TABLE OF CONTENTS Page ACKNCMLEDGEMENTS ii LIST OF FIGURES iv ABSTRACT v INTRODUCTION METHODS STUDY AREA 4 RESULTS AND DISCUSSION 4 LITERATURE CITED 14 APPENDIX A 16 VITA 25 iv LIST OF FIGURES Figure Page l. Leaf photosynthesis rates and leaf RGR of Geum ross ii in the Kob res ia site and the Des cha� a site. Each point on the photosynthesis figure is the mean of four to eight rep! icates with the vertical bars representing+ one standard error. Each point on the leaf RGR figure is the mean of ten plants with the standard errors ranging from 0.01 to 0.58. Negative leaf RGR denotes leaf senescence 5 2. Leaf photosynthesis rates and leaf RGR of Kobresia myosuroides in the Kobresia site and Deschampsia caespitosa in the Deschampsia site. Each point on the photosynthesis figure is the mean of four to eight replicates with the vertical bars repre senting+ one standard error. Each point on the leaf RGR-figure is the mean of ten plants with the standard errors ranging from 0.01 to 0.94. Negative leaf RGR denotes leaf senescence . 6 3. Relative photosynthetic contributions of Deschampsia caespitosa, Geum rossi i, and Kobresia myosuroides at different leaf positions at three dates during the growing season. Each leaf position represents the mean of four to eight rep! icates 10 v ABSTRACT Field Measurements of Photosynthesis and Leaf Growth Rates of Three Alpine Plant Species by Douglas A. Johnson, Master of Science Utah State University, 1973 Ma jor Professor: Martyn M. Caldwell Department: Range Science 14 Leaf photosynthetic measurements using a portable co field 2 system were carried out and correlative leaf relative growth rates, RGR, were determined at different leaf positions of three alpine pl ant species throughout the growing season. Initially there was a period of high leaf RGR associated with a period of increasing photosynthetic activity. Following this stage wa s a long period of no net change in length of the living leaf. During this period, photosynthetic activity generally increased to a maximum level and then decreased steadily. The final ontogenetic stage was a period of negative leaf RGR denoting leaf senescence which was associated with a marked dee] ine in leaf co uptake. Ontogenetic timing of these 2 alpine species is geared with the surge and dee] ine of individual le af photosynthetic activity so that one to several leaves operating at near ma ximal photosynthetic capacity are always maintained during the growing season for each plant, These findings are discussed in relation to their adaptive significance for these species. 00 pages) INTRODUCTION Photosynthesis of individual leaves exhibits an increase after leaf emergence, a leveling off before or after full leaf expansion, and a dee! ine with leaf senescence. This pattern has become apparent from field co uptake studies done for individual leaves of agricultural 2 species (e.g., Beuerlein and Pendleton, 1971; Turner and lncoll, 1971). In alpine areas where the growing season is short and relatively severe field photosynthesis measurements of whole plants have been carried out (e.g., Scott and Billings, 1964; Moser, 1973). However, little or no field work has been done in alpine areas with photosynthesis of individual leaves. The present investigation with three major alpine tundra species was concerned with investigating the field photo- synthetic response of individual leaves at several leaf positions and correlating this photosynthetic response with the seasonal progressions of leaf growth rate during the alpine growing season. METHODS Leaf photosynthetic measurements were determined using a 14 portable co field system. The field system and procedure have 2 been described in detail by Tieszen, et al. (in press). The field system consists of a plexiglass leaf chamber which is temperature controlled by a Peltier thermoelectric stage. The procedure involves 14 exposing an intact leaf to a co air mixture, immediately cooling the leaf sample in the field following2 exposure, subsequent drying of the exposed leaf sample, combusting this sample, and radioactivity counting using I iquid scintillation techniques. 2 14 According to studies by Ludwig and Canvin (1971) net co 2 uptake in a leaf is maximum at exposure periods less than 30 sec. This initial period approximates what might be termed as gross co 2 influx. After 30 sec, net co uptake decreases as the exposure time 2 14 in co is extended and after 10 min approaches normal net co 2 2 14 exchange rates. This decrease in co uptake is associated with 2 14 co evolution from the leaf. Thus, with exposure times of min 2 used in this study, co uptake measured was between gross co uptake 2 2 and net photosynthesis. All co uptake determinations were carried out under a constant 2 chamber temperature of 10.0 �.soc which is representative of the mean daytime growing season temperatures at the study area. These were measured using a shielded copper-constantan fine wire thermocouple. Leaf temperatures were also monitored and were always within ±1 .o0c of chamber air temperature. Artificial irradiation was provided by a high intensity incandescent lamp and was monitored with a Lambda Co. Model Ll-190SR quantum sensor. A constant intensity of 2700±200 2 l microeinsteins.m- ·sec- in the 400-700 nm wavelength range was used for all determinations and is approximately equal to maximum midday solar radiation values. A period of five minutes was allowed for the leaf to attain a steady state gas exchange rate under these 14 environmental conditions before co exposure. These temperature 2 and radiation conditions were used consistently in this study in order to assess the combined influence of other factors such as leaf age and leaf position. Leaf areas for Deschampsia caespitosa and Kobresia myosuroides were determined by direct measurement of leaf length and width and 3 are accurate to ~5%. Area values for the highly dissected leaves of Geum rossi i were estimated from regression relationships of leaf dry weight, leaf length and width, and leaf area which was determined by a modified optical planimeter (Caldwell and Moore, 1971). These basic relationships were determined at three times during the season. For the leaf growth rate study, ten plants each of Geum rossii and Kobresia myosuroides in the Kobresia meadow site and ten plants each of Deschampsia caespitosa and Geum rossii in the Deschampsia meadow site were selected on 15 June. These individual plants were observed and measured every five days for leaf emergence, leaf elongation, and senescence throughout the course of the growing season. Leaf length was measured from the base of the plant to the most distal green portion of the outstretched leaf. Each leaf was individually measured and the same leaf was followed through emergence, expansion, and senescence. From these individual leaf measurements, the mean leaf relative growth rate, RGR, was calculated by (Kvet, et al., 1971): -1 -1 RGR l e n g t h · l e n g t h · t i me ( l ) where 1 and 1 are the lengths at times t and t respectively. 2 1 2 1 In both the leaf photosynthetic and lea f RGR studies the same numerical id e ntification of leaf position was used. The first green leaf present on the plant and thus the oldest living leaf on the plant was given the designation of # 1 leaf, followed by the second green leaf to emerge as #2 leaf, with successive numbering up to the youngest leaf. 4 STUDYAREA The area of the study was on Niwot Ridge (40° 04 1 N, 1050 36 1 N) at an elevation of 3476 m in the Front Range of the Colorado Rocky Mountains. The study sites were the U.S. Tundra Biome intensive sites, one a Kobresia community and the other a Deschampsia meadow. The Kobresia site has a southwest aspect and 5° slope, while the Deschampsia site has a southeast aspect and a 4° slope.