Shoot structure and photosynthetic efficiency along the light gradient in a Scots pine canopy.

We examined the effects of structural and physiological acclimation on the photosynthetic efficiency of Scots pine (Pinus sylvestris L.) shoots. We estimated daily light interception (DLI) and photosynthesis (DPHOT) of a number of sample shoots situated at different positions in the canopy. Photosynthetic efficiency (epsilon) was defined as the ratio of DPHOT to the potential daily light interception (DLI(ref)) defined as the photosynthetically active radiation (PAR) intercepted per unit area of a sphere at the shoot location. To calculate DLI(ref), DLI and DPHOT, the radiation field surrounding a shoot in the canopy was first modeled using simulated directional distributions of incoming PAR on a clear and an overcast day, and estimates of canopy gap fraction in different directions provided by hemispherical photographs. A model of shoot geometry and measured data on shoot structure and photosynthetic parameters were used to simulate the distribution of PAR irradiance on the needle surface area of the shoot. Photosynthetic efficiency (epsilon) was separated into light-interception efficiency (epsilon(I) = DLI/DLI(ref)) and conversion efficiency (epsilon(PHOT) = DPHOT/DLI). This allowed us to quantify separately the effect of structural acclimation on the efficiency of photosynthetic light capture (epsilon(l)), and the effect of physiological acclimation on conversion efficiency (epsilon(PHOT)). The value of epsilon increased from the top to the bottom of the canopy. The increase was largely explained by structural acclimation (higher epsilon(I)) of the shade shoots. The value of epsilon(PHOT) of shade foliage was similar to that of sun foliage. Given these efficiencies, the clear-day value of DPHOT for a sun shoot transferred to shade was only half that of a shade shoot at its original position. The method presented here provides a tool for quantitatively estimating the role of acclimation in total canopy photosynthesis.

Effects of light on shoot geometry and needle morphology in Abies amabilis.

In some conifers, shoot geometry and needle morphology vary significantly in response to the light conditions under which they develop. We measured shoot length, silhouette area, total projected needle area, total needle weight and needle thickness on current shoots developed under a wide range of light conditions in a 36-year-old Abies amabilis (Dougl.) Forbes stand. Current light was quantified by evaluating percent openness from hemispherical photographs taken before the growing season. Unweighted total openness was correlated with shoot geometry and needle morphology better than any weighted indices tested. Needle thickness and leaf mass/area were both closely correlated with total openness (R(2) = 0.86 and 0.82, respectively). The most exposed needles were 2.5 times thicker and had 3-4 times more leaf mass/area than the most shaded needles. Total projected leaf area/shoot silhouette area was also correlated with openness (R(2) = 0.74) and was about twice as high in sun shoots as in shaded shoots. As a result of greater leaf mass/leaf area and greater leaf area/shoot silhouette area, a unit of intercepted light was dispersed over about 6 times as much leaf mass in a sun shoot as in a shade shoot, which presumably permits more efficient utilization of the intercepted light under high illumination with less energy wastage to light saturation. Moreover, leaf mass per unit of silhouette area was almost exactly proportional to canopy openness, as predicted by resource optimization theory if nitrogen concentration and photosynthetic capacity per unit mass are constant in new leaves. The close correlation of needle thickness and leaf mass/area with openness suggests that either parameter could be used as an index of the distribution of light or light-driven processes in an A. amabilis canopy.

Acclimation responses of mature Abies amabilis sun foliage to shading.

This paper addresses two main questions. First, can evergreen foliage that has been structurally determined as sun foliage acclimate physiologically when it is shaded? Second, is this acclimation independent of the foliage ageing process and source-sink relations? To investigate these questions, a shading and debudding experiment was established using paired branches on opengrown Abies amabilis trees. For each tree, one branch was either shaded, debudded, or both, from before budbreak until the end of summer, while the other branch functioned as a control. Foliage samples were measured both prior to and during treatment for photosynthesis at light saturation (A max), dark respiration, nitrogen content, chlorophyll content, chlorophyll-to-nitrogen ratio and chlorophyll a:b ratio. All age classes of foliage responded similarly during the treatment, although pre-treatment values differed between age classes. Within 1 month after the treatment began, A max was lower in shaded foliage and remained lower throughout the treatment period. For debudded branches, A max was lower than the controls only during active shoot elongation. At the end of the treatments in September, A max in shade-treated sun foliage matched the rates in the true shade-formed foliage, but nitrogen remained significantly higher. By 1.5 months after treatment, chlorophyll content in shaded foliage was higher than in controls, and the chlorophyll a:b ratio was lower for the shaded foliage. On debudded branches, chlorophyll content and chlorophyll a:b ratio were similar to the values in control samples. Shading lowered the rate of nitrogen accumulation within a branch, while removing debudding decreased the amount of sequestered N that was exported from the older foliage to supply new growth. By September, chlorophyll content in shade-treated foliage was higher than that in the control sun foliage or in true shade foliage. The chlorophyll increase as a result of shading was unexpected. However, the chlorophyll-to-nitrogen ratio was identical for the shade-treated sun foliage and the true shade foliage while being significantly lower than the control sun foliage. It appears that acclimation to shading in mature foliage involves a reallocation of nitrogen within the leaf into thylakoid proteins. A redistribution of resources (nitrogen) among leaves is secondary and appears to function on a slower time scale than reallocation within the leaf. Thus, A. amabilis foliage that is structurally determined as sun foliage can acclimate to shade within a few months; this process is most likely independent of ageing and is only slightly affected by source-sink relations within a branch.

Foliage dark respiration in Abies amabilis (Dougl.) Forbes: variation within the canopy.

Dark respiration of foliage was measured in a 30-year-old stand of Abies amabilis in western Washington from June to November. Both laboratory and field measurements were used to study the effect of environmental and tree variables on respiration. Foliage respiration rates were most strongly influenced by needle temperature. After accounting for leaf temperature differences, foliage respiration decreased with depth in the canopy for all age classes of foliage. Respiration differences attributed to location within the canopy were greatest early in the growing season, but were still significant in November. Younger foliage respired more than older foliage in the upper canopy, but not in the lower canopy. Respiration differences due to foliage age were highly significant in the early growing season, but were not detectable by mid-October.

Natural disturbance and the steady state in high-altitude balsam fir forests.

Wind-induced, cyclic waves of death, regeneration, and maturation constantly move through the high-altitude balsam fir forests in the northeastern United States. Biomass and productivity relations, species diversity, and nutrient cycling patterns are closely tied to this cycle of disturbance. Disturbance is thus an integral part of the long-term maintenance of this ecosystem. Since forests of this type normally include all phases of the disturbance-regeneration cycle, they may constitute a steady-state ecosystem in equilibrium with the surrounding environment.