The contribution of changes in meristem behavior to age-related decline in forest productivity is poorly understood. We studied age-related trends in needle morphology and gas exchange in a population of red spruce (Picea rubens Sarg.) growing in a multi-cohort stand where trees ranged from first-year germinants to trees over 150 years old, as well as in grafted scions from these trees. In the field study, age-related trends in foliar morphology were determined in six cohorts ranging in age from 2 to 120 years, and differences in gas exchange characteristics were compared between 60- and 120-year age classes. In a common-rootstock study, scions from trees representing 20-, 60-, and 120-year cohorts were grafted onto juvenile rootstock and maintained for three growing seasons, after which morphological and physiological foliar attributes were evaluated. The field study revealed significant age-related trends in foliar morphology, including decreasing specific leaf area, and increasing needle width, projected area, and width/length ratio. Similar trends were apparent in foliage from the grafted scions. Both in situ foliage and shoots of grafted scions from the oldest cohort showed significantly lower photosynthetic rates than their counterparts from younger trees; however, differences in stomatal conductance and internal CO(2) concentrations were not significant. These results suggest that: (1) foliage of red spruce exhibits age-related trends in both morphology and physiology; (2) age-related decreases in photosynthetic rates contribute to declining productivity in old red spruce; (3) declines in photosynthetic rates result from nonstomatal limitations; and (4) age-related changes in morphology and physiology are inherent in meristems and persist for at least 3 years in scions grafted to juvenile rootstock.

We examined the dynamics of down coarse woody debris (CWD) under an expanding-gap harvesting system in the Acadian forest of Maine. Gap harvesting treatments included 20% basal area removal, 10% basal area removal, and a control. We compared volume, biomass, diameter-class, and decay-class distributions of CWD in permanent plots before and 3 years after harvest. We also determined wood density and moisture content by species and decay class. Mean pre-harvest CWD volume was 108.9 m3/ha, and biomass was 23.22 Mg/ha. Both harvesting treatments increased the volume and biomass of non-decayed, small-diameter CWD (i.e., logging slash), with the 20% treatment showing a greater increase than the 10% treatment and both treatments showing greater increases than the control. Post-harvest reduction of advanced-decay CWD due to mechanical crushing was not evident. A mean of 18.48 m3 water/ha (1.85 L/m2) demonstrates substantial water storage in CWD, even during an exceptionally dry sampling period. The U-shaped temporal trend in CWD volume or biomass seen in even-aged stands may not apply to these uneven-aged stands; here, the trend is likely more complex because of the superimposition of small-scale natural disturbances and repeated silvicultural entries.

Woody plants exhibit significant and predictable patterns of change in morphology and physiology as they become older and larger. Four models of potential pathways controlling these changes are presented: a stimulus-response model in which fully developed organs respond to changes in environment (defined here as everything external to the organ); an extrinsic model in which the attributes of developing organs are determined by environmental factors; an intrinsic model in which changes are a result of programmed changes in gene expression; and an extrinsic-intrinsic model in which changes in gene expression are induced by environmental factors. We review evidence that a genetic component is involved in controlling age- and size-related changes in foliar morphology and physiology and discuss the possibility of complex interactions among model pathways.

The roles of temperature (T) and leaf-to-air vapor pressure deficit (VPD) in regulating net photosynthesis (A(net)) and stomatal conductance (G(s)) of red spruce (Picea rubens Sarg.) were investigated in a field study and in a controlled environment experiment. Both A(net) and G(s) exhibited a relatively flat response to temperatures between 16 and 32 degrees C. Temperatures between 32 and 36 degrees C markedly decreased both A(net) and G(s). Vapor pressure deficits above 2 kPa had significant effects on both A(net) and G(s). The influence of VPD on A(net) and G(s) fit a linear response model and did not interact significantly with T effects.

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