A field study of seasonal changes in the photosynthetic characteristics of three altitudinal populations of the snow gum (Eucalyptus pauciflora Sieb. ex. Spreng) was conducted at elevations of 915, 1215 and 1645 m in the Snowy Mountains area of south-eastern Australia. At optimal temperatures in midsummer, peak rates of photosynthesis under CO2-saturating conditions (Psub) reached levels of 125-135 ng cm-2 sec-1, and under ambient CO2 conditions (Pamb) reached levels of 45-55 ng cm-2 sec-1. Corresponding values of the intracellular resistance to CO2 transfer (ri) were 5-6 sec cm-1, and of the gas phase resistance to water-vapour transfer (r,) 1.2-2 sec cm-1. Measured at temperatures of 15, 20 and 25°C, the peak seasonal values of Pmax showed no significant difference between sites, but at 5° and 10° peak values were highest at the highest elevation (coldest site), and at 30° and 35° peak values were highest at the lowest elevation (warmest site). Generally similar patterns applied to Paamb. These features are consistent with the view that E. Pauciflora shows continuous variation in many physiological characteristics over its altitudinal range. At each site the temperature optimum for photosynthesis changed markedly during the season, and was closely correlated both with the long-term maximum air temperature and with the mean maximum temperature of the 10 days prior to the date of measurement. This appeared to reflect long-term adaptation of the photosynthetic apparatus of each population to the general temperature conditions at each site, combined with a short-term acclimation to the prevailing seasonal temperature regime.
Photosynthetic temperature response curves were measured at leaf temperatures from 10 to 40°C on seedlings of E. pauciflora grown from seed collected at tree-line (elevation 1905 m) in the Snowy Mountains area and at three lower elevations, 915, 1215 and 1645 m, which correspond to those used in an earlier field study (Slatyer and Morrow 1977). The material was grown in naturally lit, temperature-controlled greenhouses at day/night temperatures of 8/4, 15/10, 21/16, 27/22 and 33/28°C. Comprehensive measurements were made on the tree-line population, in which peak rates of net photosynthesis, Pamb, reached 75 ng cm-2 s-1 at a temperature of 20�°C, from material grown at 21/16°. Minimum levels of intracellular resistance, rt, were 2.8 s cm-1, and of leaf gas-phase resistance to CO2 transfer, r1, were 3.2 s cm-1. Changes in rt and r1, with measurement temperature, appeared to be of approximately equal importance in mediating the overall photosynthetic temperature response. Changes in the CO2 compensation point, Γ were of increasing importance at higher measurement temperatures. The photosynthetic temperature optimum was markedly affected by the growth temperature regime. In the tree-line population, it increased from about 16° when grown at 8/4° to 24° when grown at 33/28°. The relationship between the observed photosynthetic temperature optimum and the day temperature of the growth regime indicated a preferred temperature for photosynthesis of 20.0°, and a tendency for the temperature optimum to shift by 0.34° per degree shift in the day growth temperature. A similar effect of growth temperature on the photosynthetic temperature optimum was noted in the three lower-elevation populations, in which preferred temperatures of 21.5, 24.2 and 27.2° were calculated for the material collected at 1645, 1215 and 915 m respectively. These temperatures were several degrees higher than the field-observed temperature optima, although the gradient of preferred temperature with elevation was comparable to that noted in the field study.
Established, field-grown, seedlings of Eucalyptus pauciflora were transferred from high- and low- elevation field sites to a controlled-environment greenhouse in Canberra (maximum/minimum daily temperature range 26/15°) and the pattern of photosynthetic acclimation observed. Levels of net photosynthesis, Pamb, intracellular resistance rI, and leaf gas-phase resistance to CO2 transfer (r1) were monitored, as were the temperature optima for these parameters. Acclimation proceeded most rapidly in the material grown at the warmer, low-elevation, site (955 m), and in the low-elevation population. Daily maximum/minimum temperatures at this site for the 10 days prior to transfer averaged 23/11°. With this material, levels of, and the temperature optimum for, Pamb reached control levels within 6 days of transfer from the field environment. By comparison, Pamb in the high-elevation population grown at the high-elevation (tree-line) site (1910 m) where the 10-day temperature averaged 15/7°, did not reach control levels until 14 days after transfer, and the temperature optimum for Pamb required 20 days to reach the control level. In general, the patterns of change in rI and r1 paralleled those in Pamb. Both the level of physiological activity in the field, and the temperature differences between the field and greenhouse environment, appeared to affect the rate of acclimation. Immediately after transfer from the field, the temperature optima of the high-and low-elevation populations were close to the daily maximum temperature of the respective field environments. The temperature optimum of the high-elevation material grown at the low-elevation site was intermediate in value. At the conclusion of the acclimation period, the temperature optima of both high-elevation populations had converged to a value similar to that of the high-elevation control (about 22°); similarly, the temperature optimum of the low-elevation populations had reached the level of the low-elevation control (27°) These various temperature optima are interpreted on the basis that each population has a 'preferred' temperature which can be modified by different effective growth temperatures to yield different optima in different thermal environments. In the field, the effective temperature appears to be intermediate between the prevailing maximum and minimum temperatures.
A procedure for estimating field photosynthetic temperature optima from phytotron temperature response data, for elevational populations of E. pauciflora, is developed. It utilizes the principle that each population has a preferred temperature, Tpref, and an acclimation coefficient, α, which can be determined from phytotron-derived temperature response curves, and which enable the photosynthetic temperature optimum observed in a particular field temperature regime (Test) to be estimated from the expression Test = Tpref - α(Tpref - Tequiv), where Tequiv is a field temperature equivalent, in terms of its effect on the photosynthetic temperature optimum, to a known phytotron growth temperature. Application of the procedure to sets of field and greenhouse data suggests that when Tpref and α are based on phytotron day growth temperatures, and when Tequiv is based on the proposition that a square-wave conversion of the field day-time temperature curve is equivalent to the phytotron day growth temperature, estimates of field and greenhouse temperature optima can be made which give good agreement with observed values. The agreement is best when active, current-year tissue is used as a basis of the field observations and when single leaves rather than shoots are used for field measurements. The procedure is also used to compare actual rates of net photosynthesis, Pamb, obtained from field and phytotron studies, when both are plotted against equivalent temperature. Using this procedure, the large apparent differences between rates of net photosynthesis observed in the field and in the phytotron can be considerably reduced. This suggests that the notion of equivalent temperature may provide a useful means of minimizing the effects of physical, temperature-related differences in comparing field and phytotron responses, thereby widening the range of practical applications of phytotron experiments.
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