Photosynthetic capacity of red spruce during winter.

We measured the photosynthetic capacity (P(max)) of plantation-grown red spruce (Picea rubens Sarg.) during two winter seasons (1993-94 and 1994-95) and monitored field photosynthesis of these trees during one winter (1993-94). We also measured P(max) for mature montane trees from January through May 1995. Changes in P(max) and field photosynthesis closely paralleled seasonal changes in outdoor air temperature. However, during thaw periods, field photosynthesis was closely correlated with multiple-day temperature regimes, whereas P(max) was closely correlated with single-day fluctuations in temperature. There was a strong association between short-term changes in ambient temperature and P(max) during the extended thaw of January 1995. Significant increases in P(max) occurred within two days of the start of this thaw. Repeated measurements of cut shoots kept indoors indicated that temperature-induced increases in P(max) can occur within 3 h. Although significant correlations between P(max) and stomatal conductance (g(s)) or intracellular CO(2) concentration (C(i)) raised the possibility that increases in P(max) resulted from increases in stomatal aperture, fluctuations in g(s) or C(i) explained little of the overall variation in P(max). Following both natural and simulated thaws, P(max) increased considerably but plateaued at only 37% of the mean photosynthetic rate reported for red spruce during the growing season. Thus, even though shoots were provided with near-optimal environmental conditions, and despite thaw-induced changes in physiology, significant limitations to winter photosynthesis remained.

Physiological changes in red spruce seedlings during a simulated winter thaw.

We evaluated net photosynthesis, respiration, leaf conductance, xylem pressure potential (XPP) and cold hardiness in red spruce (Picea rubens Sarg.) seedlings exposed to either a continuous thaw (CT) or a daytime thaw with freezing nights (FN) for 8 days during mid-winter. Physiological differences between CT and FN seedlings were evident for all measured parameters. However, the temporal expression of treatment differences varied among parameters. When compared to FN seedlings, CT seedlings had higher rates of respiration following 24 h of treatment, and a higher net photosynthetic rate, leaf conductance and XPP after 48 h of treatment. The CT seedlings were significantly less cold tolerant than the FN seedlings following 4 days of thaw, whereas FN seedlings did not deharden over the 8 days of treatment. Examination of temporal trends among thaw-associated changes in physiology suggested that, although greater carbon exchange occurred as stomatal conductance increased, the transition from negative to positive net photosynthesis was not the result of increases in conductance, but may have been associated with thaw-induced increases in XPP. Because thaw-associated changes in gas exchange and cold hardiness were offset in time, we conclude that, if changes in these processes are physiologically linked, the linkage is indirect.

Winter photosynthesis of red spruce from three Vermont seed sources.

We evaluated winter (January through March) carbon assimilation of red spruce (Picea rubens Sarg.) from three Vermont seed sources grown in a common garden in northwestern Vermont. Although CO(2) exchange rates were generally low, net photosynthetic rates increased during two prolonged thaws. Significant correlations between CO(2) exchange rates and multiday air temperature means supported our observations of enhanced gas exchange during extended periods of elevated temperature. Increases in photosynthesis during thaws occurred before observed increases in leaf conductance, indicating that initial changes in photosynthesis were probably not associated with changes in stomatal aperture. Results of correlations between photosynthetic rates and PAR suggested that solar irradiance did not have a strong effect on winter carbon capture. Rates of net photosynthesis differed among seed sources. Trees from the Mt. Mansfield source had the highest average rates of photosynthesis and, at times, rates for individual trees from this source approximated those occurring during the growing season. Because seed sources differed in photosynthetic rates but not in leaf conductance, we conclude that differences in winter photosynthesis among seed sources were primarily attributable to factors other than changes in stomatal aperture.