Understanding and predicting ecosystem functioning (e.g., carbon and water fluxes) and the role of soils in carbon storage requires an accurate assessment of plant rooting distributions. Here, in a comprehensive literature synthesis, we analyze rooting patterns for terrestrial biomes and compare distributions for various plant functional groups. We compiled a database of 250 root studies, subdividing suitable results into 11 biomes, and fitted the depth coefficient β to the data for each biome (Gale and Grigal 1987). β is a simple numerical index of rooting distribution based on the asymptotic equation Y=1-β, where d = depth and Y = the proportion of roots from the surface to depth d. High values of β correspond to a greater proportion of roots with depth. Tundra, boreal forest, and temperate grasslands showed the shallowest rooting profiles (β=0.913, 0.943, and 0.943, respectively), with 80-90% of roots in the top 30 cm of soil; deserts and temperate coniferous forests showed the deepest profiles (β=0.975 and 0.976, respectively) and had only 50% of their roots in the upper 30 cm. Standing root biomass varied by over an order of magnitude across biomes, from approximately 0.2 to 5 kg m. Tropical evergreen forests had the highest root biomass (5 kg m), but other forest biomes and sclerophyllous shrublands were of similar magnitude. Root biomass for croplands, deserts, tundra and grasslands was below 1.5 kg m. Root/shoot (R/S) ratios were highest for tundra, grasslands, and cold deserts (ranging from 4 to 7); forest ecosystems and croplands had the lowest R/S ratios (approximately 0.1 to 0.5). Comparing data across biomes for plant functional groups, grasses had 44% of their roots in the top 10 cm of soil. (β=0.952), while shrubs had only 21% in the same depth increment (β=0.978). The rooting distribution of all temperate and tropical trees was β=0.970 with 26% of roots in the top 10 cm and 60% in the top 30 cm. Overall, the globally averaged root distribution for all ecosystems was β=0.966 (r =0.89) with approximately 30%, 50%, and 75% of roots in the top 10 cm, 20 cm, and 40 cm, respectively. We discuss the merits and possible shortcomings of our analysis in the context of root biomass and root functioning.

Between 8 and 6 million years ago, there was a global increase in the biomass of plants using C 4 photosynthesis as indicated by changes in the carbon isotope ratios of fossil tooth enamel in Asia, Africa, North America and South America. This abrupt and widespread increase in C 4 biomass may be related to a decrease in atmospheric CO 2 concentrations below a threshold that favoured C 3 -photosynthesizing plants. The change occurred earlier at lower latitudes, as the threshold for C 3 photosynthesis is higher at warmer temperatures.

The depth at which plants are able to grow roots has important implications for the whole ecosystem hydrological balance, as well as for carbon and nutrient cycling. Here we summarize what we know about the maximum rooting depth of species belonging to the major terrestrial biomes. We found 290 observations of maximum rooting depth in the literature which covered 253 woody and herbaceous species. Maximum rooting depth ranged from 0.3 m for some tundra species to 68 m for Boscia albitrunca in the central Kalahari; 194 species had roots at least 2 m deep, 50 species had roots at a depth of 5 m or more, and 22 species had roots as deep as 10 m or more. The average for the globe was 4.6±0.5 m. Maximum rooting depth by biome was 2.0±0.3 m for boreal forest. 2.1±0.2 m for cropland, 9.5±2.4 m for desert, 5.2±0.8 m for sclerophyllous shrubland and forest, 3.9±0.4 m for temperate coniferous forest, 2.9±0.2 m for temperate deciduous forest, 2.6±0.2 m for temperate grassland, 3.7±0.5 m for tropical deciduous forest, 7.3±2.8 m for tropical evergreen forest, 15.0±5.4 m for tropical grassland/savanna, and 0.5±0.1 m for tundra. Grouping all the species across biomes (except croplands) by three basic functional groups: trees, shrubs, and herbaceous plants, the maximum rooting depth was 7.0±1.2 m for trees, 5.1±0.8 m for shrubs, and 2.6±0.1 m for herbaceous plants. These data show that deep root habits are quite common in woody and herbaceous species across most of the terrestrial biomes, far deeper than the traditional view has held up to now. This finding has important implications for a better understanding of ecosystem function and its application in developing ecosystem models.

[1] Photosynthesis and respiration impart distinct isotopic signatures to the atmosphere that are used to constrain global carbon source/sink estimates and partition ecosystem fluxes. Increasingly, the ''Keeling plot'' method is being used to determine the carbon isotope composition of ecosystem respiration (d 13 C R ) in order to better understand the processes controlling ecosystem isotope discrimination. In this paper we synthesize emergent patterns in d 13 C R by analyzing 146 Keeling plots constructed at 33 sites across North and South America. In order to interpret results from disparate studies, we discuss the assumptions underlying the Keeling plot method and recommend standardized methods for determining d 13 C R . These include the use of regression calculations that account for error in the x variable, and constraining estimates of d 13 C R to nighttime periods. We then recalculate d 13 C R uniformly for all sites. We found a high degree of temporal and spatial variability in C 3 ecosystems, with individual observations ranging from À19.0 to À32.6%. Mean C 3 ecosystem discrimination was 18.3%. Precipitation was a major driver of both temporal and spatial variability of d 13 C R , suggesting (1) a large influence of recently fixed carbon on ecosystem respiration and (2) a significant effect of previous climatic effects on d 13 C R . These results illustrate the importance of water availability as a key control on atmospheric 13 CO 2 and highlight the potential of d 13 C R as a useful tool for integrating environmental effects on dynamic canopy and ecosystem processes.

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In the semiarid Intermountain West, boxelder, Acer negundo var. interior, a deciduous, dioecious tree, exhibits significant habitat-specific sex ratio biases. Although the overall sex ratio (male/female) does not deviate significantly from one, the sex ratio is significantly male biased (1.62) in drought-prone habitats, while it is significantly female biased (0.65) in moist, streamside habitats. The causes underlying gender-specific habitat associations in this species are not known. We hypothesized that spatial segregation of the sexes is maintained by differences in gender-specific photosynthetic behavior, water relations characteristics, and both instantaneous and integrated water-use efficiency. Genderspecific physiological characteristics were measured and related to growth, reproduction, population age structure, and habitat distribution of male and female trees.Under both field and controlled-environment conditions, males and females differed significantly in a number of physiological traits. Males maintained lower stomatal conductance to water vapor (g), transpiration (E), net carbon assimilation (A), leaf internal C0 2 concentration (c;), carbon isotope discrimination (D.; an index of time-integrated C; and water-use efficiency), and higher instantaneous (AI E) and long-term (D.) water-use efficiency than females. Furthermore, male trees exhibited greater stomatal sensitivity to both declining soil water content and increasing leaf-to-air vapor pressure gradients, a measure of evaporative demand. Higher rates of carbon fixation in female trees were correlated with higher g, higher leaf nitrogen concentrations, and greater stomatal densities. For females growing in both wet and dry habitats, vegetative shoots had higher growth rates than reproductive shoots, while for males, growth rates of the two shoot types did not differ. In streamside habitats, female trees exhibited significantly greater vegetative shoot growth when compared to male trees. In contrast, males showed slightly greater vegetative and much greater reproductive shoot growth in non-streamside habitats. Regardless of habitat or growing conditions, females allocated proportionately more of their aboveground biomass to reproduction than did males.These results suggest that ( 1) gender-specific physiological traits can help explain the maintenance ofhabitat-specific sex ratio biases in A. negundo along a soil moisture gradient, and (2) that the combination of the gender-specific physiology, growth, and allocation differences contribute to differences in the size (=age) structure of male and female plants within the population. Gender-specific physiological differences may have evolved as a product of selection to meet significantly different costs associated with reproduction in male and female plants.

There is an increasing ecological interest in understanding the gradients in H(2)(18)O enrichment in leaf water (i.e. a Péclet effect), because an appreciation of the significance of the Péclet effect is important for improving our understanding of the mechanistic processes affecting the (18)O composition of leaf water and plant organic material. In data sets where both source water and leaf water (18)O data are available, we can evaluate the potential contribution of a Péclet effect. As an example, we recalculate data published earlier by Roden and Ehleringer (1999, Oecologia 121:467-477) as enrichments in leaf water (Delta(L)) and cellulose (Delta(cell)) above source water. Based on these recalculations, we present support for the relevance of a Péclet effect in leaves. Further, we demonstrate that the subtle variations in Delta(L) and Delta(cell) caused by a Péclet effect may be masked in experimental systems in which variation in the source water oxygen isotope ratio is considerable.