Temporal variation of nonstructural carbohydrates in montane conifers: similarities and differences among developmental stages, species and environmental conditions.

Nonstructural carbohydrates (NSCs) are commonly used to assess the balance of carbon sources and sinks in plants. A notable application of this approach has been tests of hypotheses on carbon limitations of trees at their upper altitudinal limits, near the alpine. How NSCs vary in time is not well known in conifers during their critical seedling stage, despite the importance of knowing the temporal variations of NSCs to use snapshot measurements of NSCs to assess carbon balance. We measured NSCs in needles, separately as soluble sugars and starch; (1) over diurnal periods in seedlings of Pseudotsuga menziesii (Mirb.) Franco (a timberline species that does not occur up to treeline), (2) throughout the growth season in the seedlings of P. menziesii and Abies lasiocarpa (Hook.) Nutt. (a species that does occur up to treeline) growing along an elevation gradient in the timberline ecotone and furthermore (3) compared seedlings and co-occurring adults to assess variation with developmental stage. We also compared NSCs in seedlings grown under field or laboratory conditions to separate environmental from intrinsic factors affecting NSCs during early emergence. Diurnal variations in NSCs were minimal, especially when compared to seasonal variation, and were detectable mainly in relatively small midday maxima of soluble sugar concentrations. Seasonal patterns of NSCs were generally (and surprisingly) similar among field and laboratory seedlings and adults. Seasonal patterns of NSCs were dominated by progressive increases in soluble sugars until winter, and by early-season peaks in starch. Nonetheless, notable differences were detectable among ages, species and environmental conditions in (1) the timing and extent of the early-season maxima of starch and (2) the extent of the late-season maxima of soluble sugars. These differences in NSCs likely correspond with ecophysiologically relevant differences in carbon balance that could affect growth and survival of trees growing in the timberline ecotone.