James R. Ehleringer

Global vegetation change through the Miocene/Pliocene boundary

Between 8 and 6 million years ago, there was a global increase in the biomass of plants using C4 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 C4 biomass may be related to a decrease in atmospheric CO2 concentrations below a threshold that favoured C3-photosynthesizing plants. The change occurred earlier at lower latitudes, as the threshold for C3 photosynthesis is higher at warmer temperatures.

Maximum rooting depth of vegetation types at the global scale

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.

C4 photosynthesis, atmospheric CO2, and climate

Abstract The objectives of this synthesis are (1) to review the factors that influence the ecological, geographical, and palaeoecological distributions of plants possessing C4 photosynthesis and (2) to propose a hypothesis/model to explain both the distribution of C4 plants with respect to temperature and CO2 and why C4 photosynthesis is relatively uncommon in dicotyledonous plants (hereafter dicots), especially in comparison with its widespread distribution in monocotyledonous species (hereafter monocots). Our goal is to stimulate discussion of the factors controlling distributions of C4 plants today, historically, and under future elevated CO2 environments. Understanding the distributions of C3/C4 plants impacts not only primary productivity, but also the distribution, evolution, and migration of both invertebrates and vertebrates that graze on these plants. Sixteen separate studies all indicate that the current distributions of C4 monocots are tightly correlated with temperature: elevated temperatures during the growing season favor C4 monocots. In contrast, the seven studies on C4 dicot distributions suggest that a different environmental parameter, such as aridity (combination of temperature and evaporative potential), more closely describes their distributions. Differences in the temperature dependence of the quantum yield for CO2 uptake (light-use efficiency) of C3 and C4 species relate well to observed plant distributions and light-use efficiency is the only mechanism that has been proposed to explain distributional differences in C3/C4 monocots. Modeling of C3 and C4 light-use efficiencies under different combinations of atmospheric CO2 and temperature predicts that C4-dominated ecosystems should not have expanded until atmospheric CO2 concentrations reached the lower levels that are thought to have existed beginning near the end of the Miocene. At that time, palaeocarbonate and fossil data indicate a simultaneous, global expansion of C4-dominated grasslands. The C4 monocots generally have a higher quantum yield than C4 dicots and it is proposed that leaf venation patterns play a role in increasing the light-use efficiency of most C4 monocots. The reduced quantum yield of most C4 dicots is consistent with their rarity, and it is suggested that C4 dicots may not have been selected until CO2 concentrations reached their lowest levels during glacial maxima in the Quaternary. Given the intrinsic light-use efficiency advantage of C4 monocots, C4 dicots may have been limited in their distributions to the warmest ecosystems, saline ecosystems, and/or to highly disturbed ecosystems. All C4 plants have a significant advantage over C3 plants under low atmospheric CO2 conditions and are predicted to have expanded significantly on a global scale during full-glacial periods, especially in tropical regions. Bog and lake sediment cores as well as pedogenic carbonates support the hypothesis that C4 ecosystems were more extensive during the last glacial maximum and then decreased in abundance following deglaciation as atmospheric CO2 levels increased.

Carbon isotope ratios in belowground carbon cycle processes.

Analyses of carbon isotope ratios (δ13C) in soil organic matter (SOM) and soil respired CO2 provide insights into dynamics of the carbon cycle. δ13C analyses do not provide direct measures of soil CO2 efflux rates but are useful as a constraint in carbon cycle models. In many cases, δ13C analyses allow the identification of components of soil CO2 efflux as well as the relative contribution of soil to overall ecosystem CO2 fluxes. δ13C values provide a unique tool for quantifying historical shifts between C3 and C4 ecosystems over decadal to millennial time scales, which are relevant to climate change and land-use change issues. We identify the need to distinguish between δ13C analyses of SOM and those of soil CO2 efflux in carbon cycle studies, because time lags in the turnover rates of different soil carbon components can result in fluxes and stocks that differ in isotopic composition (disequilibrium effect). We suggest that the frequently observed progressive δ13C enrichment of SOM may be related to a g...

A mechanistic model for interpretation of hydrogen and oxygen isotope ratios in tree-ring cellulose

A mechanistic model is presented to quantify both the physical and biochemical fractionation events associated with hydrogen and oxygen isotope ratios in tree-ring cellulose. The model predicts the isotope ratios of tree-rings, incorporating both humidity and source water environmental information. Components of the model include (1) hydrogen and oxygen isotope effects associated with leaf water enrichment; (2) incorporation of leaf water isotope ratio values into photosynthetic carbohydrates along with the biochemical fractionation associated with autotrophic synthesis; (3) transport of exported carbohydrates (such as sucrose) from leaves to developing xylem in shoots and stems where cellulose is formed; (4) a partial exchange of oxygen and hydrogen isotopes in carbohydrates with xylem sap water during conversion into cellulose; and (5) a biochemical fractionation associated with cellulose synthesis. A modified version of the Craig-Gordon model for evaporative enrichment adequately described leaf water dD and d 18 O values. The leaf water model was robust over a wide range of leaf waters for both controlled experiments and field studies, far exceeding the range of values to be expected under natural conditions. The isotopic composition of cellulose was modeled using heterotrophic and autotrophic fractionation factors from the literature as well as the experimentally derived proportions of H and O that undergo exchange with xylem water during cellulose synthesis in xylem cells of tree-rings. The fraction of H and O from carbohydrates that exchange with xylem sap water was estimated to be 0.36 and 0.42, respectively. The proportions were based on controlled, long-term greenhouse experiments and field studies where the variations in the dD and d 18 O of tree-ring cellulose were measured under different source water isotopic compositions. The model prediction that tree-ring cellulose contains information on environmental water source and atmospheric vapor pressure deficit (related to relative humidity) was tested under both field and greenhouse conditions. This model was compared to existing models to explain cellulose isotope ratios under a wide range of source water and humidity conditions. Predictions from our model were consistent with observations, whereas other models showed large discrepancies as soon as the isotope ratios of source water and atmospheric water deviated from each other. Our model resolves the apparently conflicting and disparate interpretations of several previous cellulose stable isotope ratio studies. Copyright

The application and interpretation of Keeling plots in terrestrial carbon cycle research

[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 CR) in order to better understand the processes controlling ecosystem isotope discrimination. In this paper we synthesize emergent patterns in d 13 CR 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 CR. These include the use of regression calculations that account for error in the x variable, and constraining estimates of d 13 CR to nighttime periods. We then recalculate d 13 CR uniformly for all sites. We found a high degree of temporal and spatial variability in C3 ecosystems, with individual observations ranging from � 19.0 to � 32.6%. Mean C3 ecosystem discrimination was 18.3%. Precipitation was a major driver of both temporal and spatial variability of d 13 CR, suggesting (1) a large influence of recently fixed carbon on ecosystem respiration and (2) a significant effect of previous climatic effects on d 13 CR. These results illustrate the importance of water availability as a key control on atmospheric 13 CO2 and highlight the potential of d 13 CR as a useful tool for integrating environmental effects on dynamic canopy and ecosystem processes. INDEX TERMS: 0315 Atmospheric Composition and Structure: Biosphere/atmosphere interactions; 0322 Atmospheric Composition and Structure: Constituent sources and sinks; 1615 Global Change: Biogeochemical processes (4805); 1694 Global Change: Instruments and techniques; 3322 Meteorology and Atmospheric Dynamics: Land/atmosphere interactions; KEYWORDS: carbon cycle, carbon isotopes, ecosystem respiration, carbon dioxide, terrestrial ecosystems

Climate change and the evolution of C4 photosynthesis

Plants assimilate carbon by one of three photosynthetic pathways, commonly called the C(3), C(4), and CAM pathways. The C(4) photosynthetic pathway, found only among the angiosperms, represents a modification of C(3) metabolism that is most effective at low concentrations of CO(2). Today, C(4) plants are most common in hot, open ecosystems, and it is commonly felt that they evolved under these conditions. However, high light and high temperature, by themselves, are not sufficient to favor the evolution of C(4) photosynthesis at atmospheric CO(2) levels significantly above the current ambient values. A review of evidence suggests that C(4) plants evolved in response to a reduction in atmospheric CO(2) levels that began during the Cretaceous and continued until the Miocene.

Correlations between carbon isotope ratio and microhabitat in desert plants

SummaryWater is usually considered to be the key limiting factor for growth of desert plants, yet there is little information available of the water-use efficiency of species within a desert community. Leaf carbon isotope ratios, an indicator of long-term intercellular carbon dioxide concentrations and thus of water-use efficiency in C3 plants, were measured on species occurring within a Sonoran Desert community, consisting of wash, transition and slope microhabitats. Along a soil moisture gradient from the relatively wetter wash to the relatively drier slope, leaf carbon isotope ratios increased in all species, indicating that water-use efficiency increased as soil water availability decreased. Leaf carbon isotope ratios of long-lived perennials were substantially more positive than in short-lived perennials, even though plants were growing adjacent to each other. Leaf carbon isotope ratio and leaf duration (evergreen versus deciduous) were not correlated with each other. The results are discussed in terms of how the efficiency of water use may affect community structure and composition.

Differential utilization of summer rains by desert plants

SummarySeasonal changes in the hydrogen isotope ratios of xylem waters were measured to determine water sources used for growth in desert plants of southern Utah. While all species used winter-spring recharge precipitation for spring growth, utilization of summer rains was life-form dependent. Annuals and succulent perennials exhibited a complete dependence on summer precipitation. Herbaceous and woody perennial species simultaneously utilized both summer precipitation and remaining winter-spring precipitation, with herbaceous species much more reliant on the summer precipitation component. Several of the woody perennials exhibited no response to summer precipitation. Currently, precipitation in southern Utah is evenly partitioned between winter and summer time periods; however, global circulation models predict that summer precipitation will increase in response to anticipated climate change. Our data indicate that components within the community will differentially responde to the change in precipitation patterns. These results are discussed in relation to impact on competition and possible changes in community structure.

Leaf carbon isotope and mineral composition in subtropical plants along an irradiance cline

SummaryLeaf carbon isotope ratios and leaf mineral composition (Ca, K, Mg, Mn, N, and P) were measured on the dominant species along an irradiance cline in a subtropical monsoon forest of southern China. This irradiance cline resulted from disturbance caused by fuel-harvesting. Leaf carbon isotope ratios increased from undisturbed to disturbed sites for all species, indicating that leaf intercellular CO2 concentrations decreased and leaf water use efficiencies increased along this cline. Nitrogen and magnesium levels were lower in leaves of species on the disturbed sites, but there were no clear patterns for calcium, potassium, phosphorus or manganese.