The scaling of respiratory metabolism with body mass is one of the most pervasive phenomena in biology. Using a single allometric equation to characterize empirical scaling relationships and to evaluate alternative hypotheses about mechanisms has been controversial. We developed a method to directly measure respiration of 271 whole plants, spanning nine orders of magnitude in body mass, from small seedlings to large trees, and from tropical to boreal ecosystems. Our measurements include the roots, which have often been ignored. Rather than a single power-law relationship, our data are fit by a biphasic, mixed-power function. The allometric exponent varies continuously from 1 in the smallest plants to 3/4 in larger saplings and trees. The transition from linear to 3/4-power scaling may indicate fundamental physical and physiological constraints on the allocation of plant biomass between photosynthetic and nonphotosynthetic organs over the course of ontogenetic plant growth.allometry | metabolic scaling | mixed-power function | whole-plant respiration | simple-power function F rom the smallest seedlings to giant trees, the masses of vascular plants span 12 orders of magnitude in mass (1). The growth rates of most plants, which are generally presented in terms of net assimilation rates of CO 2 , are believed to be controlled by respiration (2, 3). Furthermore, many of the CO 2 -budget models of plant growth and carbon dynamics in terrestrial ecosystems are based on whole-plant respiration rates in relation to plant size (2, 4-7). Thus far, however, there have been few studies of wholeplant respiration over the entire range of plant size from tiny seedlings to large trees. The purpose of the present study was to quantify the allometric scaling of metabolism by directly measuring whole-plant respiration over a representative range of sizes.For the past century, the scaling of metabolic rate with body size has usually been described using an allometric equation, or simple power function, for the form (8-17)where Y is the respiratory metabolic rate (μmol s −1 ), F is a constant (μmol s −1 kg -f ), M is the body mass (kg), and f is the scaling exponent. The exponent f has been controversial, and various values have been reported based on studies of both animals and plants (15). Recently, it was suggested that f = 1 for relatively small plants, based on data for a 10 6 -fold range of body mass (16), including measurements using a whole-plant chamber (18,19). If f = 1, this means that whole-plant respiration scales isometrically with body mass, which may be reasonable in the case of herbaceous plants and small trees because nearly all of their cells, even those in the stems, should be active in respiration. However, it was suggested that f = 3/4 based originally on empirical studies of animal metabolism (8). This idea is consistent with the mechanistic models of resource distribution in vascular systems (10, 11), including the pipe model (20, 21) and models based on space-filling, hierarchical, fractal-like networks of br...

In the temperate zone of Japan, Pinus densiflora Sieb. et Zucc. bears needles of up to three age classes in the upper crown and up to five age classes in the lower crown. To elucidate the effects of leaf age on photosynthetic parameters and its relationships with leaf mass per unit area (LMA) and leaf nitrogen (N(l)) concentration on an area (N(a)) and mass (N(m)) basis, we measured seasonal variations in LMA, N(l), light-saturated photosynthetic rate (A(max)), stomatal conductance (g(s)), maximum rate of carboxylation (V(cmax)) and maximum rate of electron transport (J(max)) in leaves of all age classes in the upper and lower crown. Leaf mass per unit area increased by 27% with increasing leaf age in the lower crown, but LMA did not depend on age in the upper crown. Leaf age had a significant effect on N(m) but not on N(a) in both crown positions, indicating that decreases in N(m) resulted from dilution. Photosynthetic parameters decreased significantly with leaf age in the lower crown (39% for A(max) and 43% for V(cmax)), but the effect of leaf age was not as great in the upper crown, although these parameters exhibited seasonal variation in both crown positions. Regression analysis indicated a close relationship between LMA and N(a), regardless of age class or when each age class was pooled (r(2) = 0.57-0.86). Relationships between LMA and N(a) and among A(max), V(cmax) and J(max) were weak or not significant when all age classes were examined by regression analysis. However, compared with older leaves, relationships among LMA, N(a) and A(max) were stronger in younger leaves. These results indicate that changes in LMA and N(l) mainly reflect light acclimation during leaf development, but they are only slightly affected by irradiance in mature leaves. In conclusion, LMA and N(l) are useful parameters for estimating photosynthetic capacity, but age-related effects need to be taken into account, especially in evergreen conifers.

We measured seasonal variation in area-based nitrogen concentration (N), maximum rate of carboxylation (Vcmax) and maximum rate of electron transport (Jmax) in 1-year-old needles along four first-order branches within a Pinus densiflora Sieb. & Zucc. crown, and analyzed their relationships to growth irradiance and temperature. Each leaf light environment was expressed as a ratio of the monthly mean of daily integrated photosynthetically active irradiance (Iint) for the particular needle to Iint above the canopy (Irel). Needle N decreased in the upper crown during the development of new needles, whereas it remained fairly constant in the lower crown, reflecting differences between upper and lower crown needles in their contribution to the nitrogen of new needles. Gradients of N within the crown were correlated with Irel in all seasons (r2 = 0.40-0.78). Seasonal variation in N was weakly correlated with mean daily air minimum temperatures. Both Vcmax and Jmax showed seasonal variation in all first-order branches, and decreased to their lowest values in winter. The gradients of Vcmax and Jmax within the crown were not correlated with Irel in some seasons, but were correlated with changes in N in most months (r2 = 0.33-0.75), except in the winter. Furthermore, the regression slope of the relationship between N and Vcmax and the temperature response of Vcmax and Jmax exhibited seasonal variation.

We measured horizontal and vertical gradients of light (rPPFD) along four first-order branches of a Pinus densiflora Sieb. & Zucc. crown, and compared variations in specific leaf area (SLA), needle nitrogen concentration (N), chlorophyll concentration (Chl) and photosynthetic capacity (i.e., maximum rate of carboxylation (V(cmax))) along the two axes. The horizontal gradient of rPPFD along first-order branches was similar in magnitude to the vertical gradient of rPPFD from the upper to the lower crown. None of the measured parameters (i.e., SLA, N, Chl and Vcmax) were strictly proportional to rPPFD, although they were more or less correlated with light when data obtained for all of the crown were pooled (r(2) = 0.31-0.80). The slope of rPPFD against N on an area basis (Narea) for a branch in the middle of the crown orientated northward was significantly greater than the slope for a similar branch orientated southward. Horizontal variations were unrelated to age effects because measurements were all on 1-year-old needles. We conclude that factors other than light (i.e., orientation) may influence N allocation within branches. There was considerably less variation in the relationship of Vcmax to Narea (r2 = 0.58) than in the relationship of Vcmax to rPPFD (r2 = 0.41). Fractional N distribution among components of the photosynthetic machinery was constant within the crown. Together with the relationships between rPPFD and N on a mass basis (r2 = 0.80) and SLA and Vcmax (r2 = 0.60), these findings suggest that most light acclimation in P. densiflora occurs through changes in needle morphology (e.g., SLA) during development.

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