Root traits of perennial C4 grasses contribute to cultivar variations in soil chemistry and species patterns in particulate and mineral-associated carbon pool formation

Recent studies have indicated that the C₄ perennial bioenergy crops switchgrass (Panicum virgatum) and big bluestem (Andropogon gerardii) accumulate significant amounts of soil carbon (C) owing to their extensive root systems. Soil C accumulation is likely driven by inter- and intraspecific variability in plant traits, but the mechanisms that underpin this variability remain unresolved. In this study we evaluated how inter- and intraspecific variation in root traits of cultivars from switchgrass (Cave-in-Rock, Kanlow, Southlow) and big bluestem (Bonanza, Southlow, Suther) affected the associations of soil C accumulation across soil fractions using stable isotope techniques. Our experimental field site was established in June 2008 at Fermilab in Batavia, IL. In 2018, soil cores were collected (30 cm depth) from all cultivars. We measured root biomass, root diameter, specific root length, bulk soil C, C associated with coarse particulate organic matter (CPOM) and fine particulate organic matter plus silt- and clay-sized fractions, and characterized organic matter chemical class composition in soil using high-resolution Fourier-transform ion cyclotron resonance mass spectrometry. C₄ species were established on soils that supported C₃ grassland for 36 years before planting, which allowed us to use differences in the natural abundance of stable C isotopes to quantify C₄ plant-derived C. We found that big bluestem had 36.9% higher C₄ plant-derived C compared to switchgrass in the CPOM fraction in the 0–10 cm depth, while switchgrass had 60.7% higher C₄ plant-derived C compared to big bluestem in the clay fraction in the 10–20 cm depth. Our findings suggest that the large root system in big bluestem helps increase POM-C formation quickly, while switchgrass root structure and chemistry build a mineral-bound clay C pool through time. Thus, both species and cultivar selection can help improve bioenergy management to maximize soil carbon gains and lower CO₂ emissions.     

Determination of root biomasses of three species grown in a mixture using stable isotopes of carbon and nitrogen

A method is evaluated that employs variation in stable C and N isotopes from fractionations in C and N acquisition and growth to predict root biomasses of three plant species in mixtures. Celtis laevigata Willd. (C₃), Prosopis glandulosa Torr. (C₃, legume) and Schizachyrium scoparium (Michx.) Nash (C₄), or Gossypium hirsutum L. (C₃), Glycine max (L.) Merr. (C₃ legume), and Sorghum bicolor (L.) Moench (C₄) were grown together in separate, three-species combinations. Surface roots (0–10 cm depth) of each species from each of the two combinations were mixed in various proportions, and the relative abundances of ¹⁵N and ¹⁴N and ¹³C and ¹²C in prepared mixtures, surface roots of single species, and roots extracted from the 80-cm soil profile in which each species combination was grown were analyzed by mass spectrometry. An algebraic determination which employed the δ ¹³C, % ¹⁵N, and C and N concentrations of root subsamples of individual species accounted for more than 95% of the variance in biomass of each species in prepared mixtures with G. max, G. hirsutum, and S. bicolor. A similar analysis demonstrated species-specific differences in rooting patterns. Root biomasses of the C₄ monocots in each combination, S. scoparium and S. bicolor, were concentrated in the upper 20 cm of soil, while those of G. hirsutum and the woody P. glandulosa were largest in lower soil strata. Analyses of stable C and N isotopes can effectively be used to distinguish roots of species which differ in ratios of ¹⁵N to ¹⁴N and ¹³C to ¹²C and thus to study belowground competition between or rooting patterns of associated species with different C and N isotope signatures. The method evaluated can be extended to quantify aboveground and belowground biomasses of component species in mixtures with isotopes of other elements or element concentrations that differ consistently among plants of interest.     

Estimation of food composition of Hodotermes mossambicus (Isoptera: Hodotermitidae) based on observations and stable carbon isotope ratios

Abstract The diet of the harvester termite Hodotermes mossambicus was investigated at two sites with distinct dietary components: C₄ grasses (δ¹³C isotope values, ?13.8‰ to ?14.0‰) and C₃ plants (δ¹³C isotope values, ?25.6‰ to ?27.1‰). By comparing observations of food items carried into the colony by the termites and carbon isotope ratios of whole termites (that determined assimilated carbon), the relative proportion of the C₃ and C₄ plant food components of the termite diet was estimated. There was agreement between the observational data and stable carbon isotopic data, with grass representing approximately 93% of the diet of H. mossambicus at two study sites (urban and rural) on the South African highveld. However, when correcting for mass of food items, that is, C₃ and C₄, carried by termites, the proportion of grass (C₄) in the diet may be underestimated.     

Can C4 plants contribute to aquatic food webs of subtropical streams?

1. Recent stable isotope studies have revealed that C₄ plants play a minor role in aquatic food webs, despite their often widespread distribution and production. We compared the breakdown of C₃ (Eucalyptus) and C₄ (Saccharum and Urochloa) plant litter in a small rain forest stream and used laboratory feeding experiments to determine their potential contribution to the aquatic food web. 2. All species of litter broke down at a fast rate in the stream, although Urochloa was significantly faster than Eucalyptus and Saccharum. This was consistent with the observed higher total organic nitrogen of Urochloa compared with the other two species. 3. The breakdown of Urochloa and Saccharum was, however, not associated with shredding invertebrates, which were poorly represented in leaf packs compared with the native Eucalyptus. The composition of the invertebrate fauna in packs of Urochloa quickly diverged from that of the other two species. 4. Feeding experiments using a common shredding aquatic insect Anisocentropus kirramus showed a distinct preference for Eucalyptus over both C₄ species. Anisocentropus was observed to ingest C₄ plant litter, particularly in the absence of other choices, and faecal material collected was clearly of C₄ origin, as determined by stable isotope analysis. However, the stable carbon isotope values of the larvae did not shift away from their C₃ signature in any of the feeding trials. 5. These data suggest that shredders avoid the consumption of C₄ plants, in favour of native C₃ species that appear to be of lower food quality (based on C : N ratios). Lower rates of consumption and lack of assimilation of C₄ carbon also suggest that shredders may have a limited ability to process this material, even in the absence of alternative litter sources. Large scale clearing of forest and vegetation for C₄ crops such as sugarcane will undoubtedly have important consequences for stream ecosystem function.     

Stomatal density and aperture length in four plant species grown across a subambient CO2 gradient

Stomatal density, stomatal aperture length, area/leaf, and number of stomata/leaf were measured after the annual C₃ agronomic grasses oats (Avena sativa) and wheat (Triticum aestivum), the C, woody legume honey mesquite (Prosopis glandulosa), and the perennial C₄ grass little bluestem (Schizachyrium scoparium) were grown across a subambient carbon dioxide concentration ([CO₂]) gradient from near 200 to 350 μmol/mol in a growth chamber. The purpose was to determine if the size and density of stomata vary in response to atmospheric [CO₂] during growth, across a subambient [CO₂] range representative of the doubling that has occurred since the last ice age. Changes in stomatal density and aperture length with increasing [CO₂] were small when detected. Stomatal density decreased on adaxial flag leaf surfaces of wheat, and aperture length increased slightly with [CO₂], Leaf area and number of stomata/flag leaf increased by similar proportions with [CO₂] in two wheat cultivars. No consistent relationship between [CO₂] and stomatal density or size was detected in mesquite, oats, or little bluestem. We conclude that individual plants of these species lack the plasticity to significantly alter stomatal density and aperture length in response to increasing atmospheric [CO₂] in a single generation (annuals) or growing season (perennials).     

The effects of clipping and soil moisture on leaf and root morphology and root respiration in two temperate and two tropical grasses

Numerous studies have explored the effect of environmental conditions on a number of plant physiological and structural traits, such as photosynthetic rate, shoot versus root biomass allocation, and leaf and root morphology. In contrast, there have been a few investigations of how those conditions may influence root respiration, even though this flux can represent a major component of carbon (C) pathway in plants. In this study, we examined the response of mass-specific root respiration (μmol CO₂ g⁻¹ s⁻¹), shoot and root biomass, and leaf photosynthesis to clipping and variable soil moisture in two C₃ (Festuca idahoensis Elmer., Poa pratensis L.) and two C₄ (Andropogon greenwayi Napper, and Sporobolus kentrophyllus K. Schum.) grass species. The C₃ and C₄ grasses were collected in Yellowstone National Park, USA and the Serengeti ecosystem, Africa, respectively, where they evolved under temporally variable soil moisture conditions and were exposed to frequent, often intense grazing. We also measured the influence of clipping and soil moisture on specific leaf area (SLA), a trait associated with moisture conservation, and specific root length (SRL), a trait associated with efficiency per unit mass of soil resource uptake. Clipping did not influence any plant trait, with the exception that it reduced the root to shoot ratio (R:S) and increased SRL in P. pratensis. In contrast to the null effect of clipping on specific root respiration, reduced soil moisture lowered specific root respiration in all four species. In addition, species differed in how leaf and root structural traits responded to lower available soil moisture. P. pratensis and A. greenwayi increased SLA, by 23% and 33%, respectively, and did not alter SRL. Conversely, S. kentrophyllus increased SRL by 42% and did not alter SLA. F. idahoensis responded to lower available soil moisture by increasing both SLA and SRL by 38% and 33%, respectively. These responses were species-specific strategies that did not coincide with photosynthetic pathway (C₃/C₄) or growth form. Thus, mass-specific root respiration responded uniformly among these four grass species to clipping (no effect) and increased soil moisture stress (decline), whereas the responses of other traits (i.e., R:S ratio, SLA, SRL) to the treatments, especially moisture availability, were species-specific. Consequently, the effects of either clipping or variation in soil moisture on the C budget of these four different grasses species were driven primarily by the plasticity of R:S ratios and the structural leaf and root traits of individual species, rather than variation in the response of mass-specific root respiration.     

Occurrence and ecological characteristics of C4 dicot and Cyperaceae species in the Hungarian flora

The non-graminaceous wild flora of Hungary was screened for C₄ plants by using the stable carbon isotope ratio, the leaf anatomy and the photosynthetic carbon dioxide compensation concentration to determine the photosynthetic pathway type. On the whole, 31 C₄ species (native or naturalized) were found in the Amaranthaceae, Chenopodiaceae, Cyperaceae, Euphorbiaceae, Portulacaceae and Zygophyllaceae families. Together with the 26 C₄ grass species (Poaceae) reported earlier (Kalapos 1991), a total of 57 wild C₄ species occur in Hungary, which forms 2.6 % of the country's angiosperm flora. This figure is somewhat higher than what was expected on climatic grounds, a fact probably due to certain edaphic conditions favouring C₄ plant growth. In Hungary, the C₄ species are predominantly annuals growing in open habitats such as dry grasslands, inland saline areas, temporarily exposed riverbeds and disturbed sites. In comparison with C₃ plants, the C₄ species have higher temperature and light preferences, and their phenology lags behind that of the C₃ plants. These differences might account for C₄ plants being usually excluded from productive biotopes in Hungary, where the C₃ canopy may become closed during the growing season before C₄ plants can start their ontogenetic development. Ecological properties of C₃ and C₄ plants differ considerably in the Cyperaceae, but much less in the Chenopodianceae family. Among C₄ annuals naturalized aliens are common, most of which colonized hungary in the last two centuries. Increasing preponderance of C₄ plants is anticipated in the future as a consequence of possible climate changes and the ever increasing human impact on terrestrial vegetation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Stable carbon isotope analysis of soil organic matter illustrates vegetation change at the grassland/woodland boundary in southeastern Arizona,USA

In southeastern Arizona, Prosopis juliflora (Swartz) DC. and Quercus emoryi Torr. are the dominant woody species at grassland/woodland boundaries. The stability of the grassland/woodland boundary in this region has been questioned, although there is no direct evidence to confirm that woodland is encroaching into grassland or vice versa. We used stable carbon isotope analysis of soil organic matter to investigate the direction and magnitude of vegetation change along this ecotone. ¹³C values of soil organic matter and roots along the ecotone indicated that both dominant woody species (C₃) are recent components of former grasslands (C₄), consistent with other reports of recent increases in woody plant abundance in grasslands and savannas throughout the world. Data on root biomass and soil organic matter suggest that this increase in woody plant abundance in grasslands and savannas may increase carbon storage in these ecosystems, with implications for the global carbon cycle.     

Carbon isotope ratios and crop analyses of Arphia (Orthoptera: Acrididae) species in southeastern Wyoming Grassland

Summary Food habits of Arphia conspersa Scudder and Arphia pseudonietana (Thomas) were studied along an altitudinal transect in southeastern Wyoming shortgrass mixed prairie. Stable carbon isotope ratios indicated that diets were significantly different between study sites, between species, and between sexes. These differences were found to be primarily related to the availability of different food plants along the transect, although species with the C₃ pathway of photosynthesis were consumed in greater proportion than their availability in the habitat. The preference for C₃ species is presumably related to their higher nutritional value and digestibility, in spite of the fact that more time and energy must be spent to locate these food plants in some of the habitats studied. This study demonstrates the utility of the carbon isotope method in studying plant-animal interactions in habitats containing both C₃ and C₄ plants.     

Carbon isotope discrimination in C3-C4 intermediates

Carbon isotope discrimination in C₃–C₄ intermediates is determined by fractionations during diffusion and the biochemical fractionations occurring during CO₂ fixation. These biochemical fractionations in turn depend on the fractionation by Rubisco in the mesophyll, the amount of CO₂ fixation. These biochemical fractionations in turn depend on the fractionation by Rubisco in the mesophyll, the amount of CO₂ fixation occurring in the bundle sheath, the extent of bundle-sheath leakiness and the contribution which C₄-cycle activity makes to the CO₂ pool there. In most instances, carbon isotope discrimination in C₃–C₄ intermediates is C₃-like because only a small fraction of the total carbon fixed is fixed in the bundle sheath. In particular, this must be the case for Flaveria intermediates which initially fix substantial amounts of CO₂ into C₄-acids. In C₃–C₄ intermediates that refix photorespiratory CO₂ alone, it is possible for carbon isotope discrimination to be greater than in C₃-species, particularly at low CO₂ pressures or at high leaf temperatures. Short-term measurements of carbon isotope discrimination and gas exchange of leaves can be used to study the photosynthetic pathways of C₃-C₄ intermediates and their hybrids as has recently been done for C₃ and C₄ species.     

The use of carbon isotope ratios to evaluate legume contribution to soil enhancement in tropical pastures

Soil carbon distribution with depth, stable carbon isotope ratios in soil organic matter and their changes as a consequence of the presence of legume were studied in three 12-year-old tropical pastures (grass alone —Brachiaria decumbens (C₄), legume alone —Pueraria phaseoloides (C₃) and grass + legume) on an Oxisol in Colombia. The objective of this study was to determine the changes that occurred in the¹³C isotope composition of soil from a grass + legume pasture that was established by cultivation of a native savanna dominated by C₄ vegetation. The¹³C natural abundance technique was used to estimate the amount of soil organic carbon originating from the legume. Up to 29% of the organic carbon in soil of the grass + legume pasture was estimated to be derived from legume residues in the top 0–2-cm soil depth, which decreased to 7% at 8–10 cm depth. Improvements in soil fertility resulting from the soil organic carbon originated from legume residues were measured as increased potential rates of nitrogen mineralization and increased yields of rice in a subsequent crop after the grass + legume pasture compared with the grass-only pasture. We conclude that the¹³C natural abundance technique may help to predict the improvements in soil quality in terms of fertility resulting from the presence of a forage legume (C₃) in a predominantly C₄ grass pasture.     

Initiation of efficient C4 pathway in response to low ambient CO2 during the bloom period of a marine dinoflagellate

Dinoflagellates are important primary producers and major causative agents of harmful algal blooms in the global ocean. Despite the great ecological significance, the photosynthetic carbon acquisition by dinoflagellates is still poorly understood. The pathways of photosynthetic carbon assimilation in a marine dinoflagellate Prorocentrum donghaiense under both in situ and laboratory-simulated bloom conditions were investigated using a combination of metaproteomics, qPCR, stable carbon isotope and targeted metabolomics approaches. A rapid consumption of dissolved CO₂ to generate high biomass was observed as the bloom proceeded. The carbon assimilation genes and proteins including intracellular carbonic anhydrase 2, phosphoenolpyruvate carboxylase, phosphoenolpyruvate carboxykinase and RubisCO as well as their enzyme activities were all highly expressed at the low CO₂ level, indicating that C₄ photosynthetic pathway functioned in the blooming P. donghaiense cells. Furthermore, δ¹³C values and content of C₄ compound (malate) significantly increased with the decreasing CO₂ concentration. The transition from C₃ to C₄ pathway minimizes the internal CO₂ leakage and guarantees efficient carbon fixation at the low CO₂ level. This study demonstrates the existence of C₄ photosynthetic pathway in a marine dinoflagellate and reveals its important complementary role to assist carbon assimilation for cell proliferation during the bloom period.     

Changes in prairie vegetation under elevated carbon dioxide levels and two soil moisture regimes

Abstract. It is important to know how increasing levels of atmospheric CO₂ will affect native vegetation. The objective of this study was to determine the effect of elevated CO₂ concentrations on species composition in a tallgrass prairie kept at a high water level (730 mm of water in a 2000 mm soil profile) and a low water level (660 mm of water in 2000 mm). 16 cylindrical plastic chambers were placed on the prairie to maintain two levels of CO₂ (ambient or twice ambient) during two growing seasons in 1989 and 1990. Frequency of species was determined on 25 July 1989 and on 5 and 10 October 1990. At the beginning of the study, Poa pratensis (Kentucky bluegrass), the dominant C₃ species, had the highest frequency of 43.3%, but decreased with time. However, at the end of the experiment and under the high soil-water level, there were more P. pratensis plants in the elevated CO₂ treatment (frequency: 13.5%) than in the ambient CO₂ treatment (1.0%). Under the low soil water regime, the reverse occurred (frequencies: 3.6% and 11.0% for high and low CO₂, respectively). The frequency of major C₄ plants, Andropogon gerardii (big bluestem), A. scoparius (little bluestem) and Sorghastrum nutans (Indian grass) was not affected by CO₂. However, water did affect their frequency. Under low water, the frequency of A. gerardii decreased between 1989 and 1990. Under both soil moisture levels, the frequencies of S. nutans and A. scoparius increased. At the end of the study, Indian grass grown with high water had the highest frequency of all species on the prairie (frequency at the end of the study in October, 1990, of 44.4% and 47.4% for the high and low CO₂ levels, respectively). Unlike Indian grass, little bluestem grew better under low water conditions than under high water conditions. These results suggest that, if the climate becomes drier, A. scoparius will flourish more than S. nutans or A. gerardii, and P. pratensis may die out. Elevated CO₂ might not increase survival of C₃ plants under dry conditions, if temperatures are too high for them.     

Stable isotopes reveal seasonal competition for resources between late Pleistocene bison (Bison) and horse (Equus) from Rancho La Brea,southern California

Determining how organisms partition or compete for resources within ecosystems can reveal how communities are assembled. The Late Pleistocene deposits at Rancho La Brea are exceptionally diverse in large mammalian carnivores and herbivores, and afford a unique opportunity to study resource use and partitioning among these megafauna. Resource use was examined in bison and horses by serially sampling the stable carbon and oxygen isotope values found within tooth enamel of individual teeth of seven bison and five horses. Oxygen isotope results for both species reveal a pattern of seasonal enamel growth, while carbon isotope values reveal a more subtle seasonal pattern of dietary preferences. Both species ate a diet dominated by C₃ plants, but bison regularly incorporated C₄ plants into their diets, while horses ate C₄ plants only occasionally. Bison had greater total variation in carbon isotope values than did horses implying migration away from Rancho La Brea. Bison appear to incorporate more C₄ plants into their diets during winter, which corresponds to previous studies suggesting that Rancho La Brea, primarily surrounded by C₃ plants, was used by bison only during late spring. The examination of intra-tooth isotopic variation which reveals intra-seasonal resource use among bison and horse at Rancho La Brea highlights the utility of isotopic techniques for understanding the intricacies of ecology within and between ancient mammals.     

Plant functional types and their ecological responses to salinization in saline grasslands, Northeastern China

Photosynthetic pathways (C₃, C₄, and CAM) and morphological functional types (e.g. shrubs, high perennial grasses, short perennial graminaceous plants, annual grasses, annual forbs, perennial forbs, halophytes, and hydrophytes) were identified for the species from salinity grasslands in Northeastern China, using the data from both stable carbon isotope ratios (¹³C) and from the references published between 1993 and 2002. 150 species, in 99 genera and 37 families, were found with C₃ photosynthesis, and most of these species are dominants [e.g. Leymus chinensis (Trin.) Tzvel., Calamagrostis epigeios (L.), Suaeda corniculata (C.A. Mey.) Bunge]. 40 species in 25 genera and 8 families were identified with C₄ photosynthesis [e.g. Chloris virgata Sw., Aeluropus littoralis (Gouan) Parlat] and 1 species with CAM photosynthesis. Gramineae is the leading family with C₄ photosynthesis (27 species), Chenopodiaceae ranks the second (5 species). The significant increase of C₄ proportions with intense salinity suggested this type plant is remarkable response to the grassland salinization in the region. 191 species were classified into eight morphological functional types and the changes of most of these types (e.g. PEF, HAL, and HPG) were consistent with habitats and vegetation dynamics in the saline grassland. My findings suggest that the photosynthetic pathways, combined with morphological functional types, are efficient means for studying the linkage between species and ecosystems in this type of saline grassland in Northeastern China.This revised version was published online in March 2005 with corrections to the page numbers.     

Variations in the foliar δ13C and C3/C4 species richness in the Japanese flora of Poaceae among climates and habitat types under human activity

For 383 Poaceae species harvested over the Japanese islands and stored as herbarium specimens along several decades, we determined C₃ and C₄ types of photosynthesis from leaf stable carbon isotope ratio (δ¹³C). Then, we sought the relationships between C₄ species richness and climatic factors or habitat types. Except for the two Panicum species (P. lanuginosum and P. scoparium) having the possibility of C₃–C₄ intermediate, 227 and 154 species were classified into C₃ and C₄. The C₄ species richness increased from northern to southern islands in Japan, positively correlated with mean annual air temperature. Greater C₄ species richness in the seashore habitats, and smaller C₄ species richness in the shaded, wet and highland habitats would be related to the photosynthetic responses to local environmental factors such as irradiance level and temperature regime. No difference of leaf δ-value of C₃ Poaceae was obtained between the habitats with different soil water availability, suggesting the less importance of soil water availability on leaf water-use efficiency in C₃ Poaceae species in Japan having humid climate. Additionally, possible effects of human activity around the harvested time or site on leaf δ-value were estimated, because the habitat includes the sites with high human activity. Leaf δ-value was decreased with sampling year, and it was higher in the densely inhabited district for both C₃ and C₄. They are probably due to a historical decrease in the atmospheric δ-value via increasing human activity, and high gas emission at the districts of high human density.     

Stable isotope analysis for assessing the intercrop movement of Microplitis mediator,a larval endoparasitoid of Helicoverpa armigera

Methods to trace source habitats and movement of parasitic natural enemies in agroecosystems are limited. This study demonstrates that stable carbon isotope analysis offers a valuable new method for revealing the movement of Microplitis mediator (Haliday) (Hymenoptera: Braconidae), a larval endoparasitoid of Helicoverpa armigera Hübner (Lepidoptera: Noctuidae), between C₃ and C₄ plants. Results indicate that M. mediator with δ¹³C values of lower than −22‰ originate from a C₃ plant, whereas those with δ¹³C values of higher than −19‰ develop on a C₄ plant.     

Turnover of carbon isotopes in tail hair and breath CO<Subscript>2</Subscript> of horses fed an isotopically varied diet

Temporal stable isotope records derived from animal tissues are increasingly studied to determine dietary and climatic histories. Despite this, the turnover times governing rates of isotope equilibration in specific tissues following a dietary isotope change are poorly known. The dietary isotope changes recorded in the hair and blood bicarbonate of two adult horses in this study are found to be successfully described by a model having three exponential isotope pools. For horse tail hair, the carbon isotope response observed following a dietary change from a C₃ to a C₄ grass was consistent with a pool having a very fast turnover rate (t _1/2~0.5 days) that made up ~41% of the isotope signal, a pool with an intermediate turnover rate (t _1/2 ~4 days) that comprised ~15% of the isotope signal, and a pool with very slow turnover rate (t _1/2 ~140 days) that made up ~44% of the total isotope signal. The carbon isotope signature of horse blood bicarbonate, in contrast, had a different isotopic composition, with ~67% of the isotope signal coming from a fast turnover pool (t _1/2 0.2 days), ~17% from a pool with an intermediate turnover rate (t _1/2 ~3 days) and ~16% from a pool with a slow turnover rate (t _1/2 ~50 days). The constituent isotope pools probably correspond to one exogenous and two endogenous sources. The exogenous source equates to our fast turnover pool, and the pools with intermediate and slow turnover rates are thought to derive from the turnover of metabolically active tissues and relatively inactive tissues within the body, respectively. It seems that a greater proportion of the amino acids available for hair synthesis come from endogenous sources compared to the compounds undergoing cellular catabolism in the body. Consequently, the isotope composition of blood bicarbonate appears to be much more responsive to dietary isotope changes, whereas the amino acids in the blood exhibit considerable isotopic inertia.     

Kangaroo tooth enamel oxygen and carbon isotope variation on a latitudinal transect in southern Australia: implications for palaeoenvironmental reconstruction

Tooth enamel apatite carbonate carbon and oxygen isotope ratios of modern kangaroos (Macropus spp.) collected on a 900-km latitudinal transect spanning a C₃–C₄ transition zone were analysed to create a reference set for palaeoenvironmental reconstruction in southern Australia. The carbon isotope composition of enamel carbonate reflects the proportional intake of C₃ and C₄ vegetation, and its oxygen isotope composition reflects that of ingested water. Tooth enamel forms incrementally, recording dietary and environmental changes during mineralisation. Analyses show only weak correlations between climate records and latitudinal changes in δ¹³C and δ¹⁸O. No species achieved the δ¹³C values (~?1.0 ‰) expected for 100 % C₄ grazing diets; kangaroos at low latitudes that are classified as feeding primarily on C₄ grasses (grazers) have δ¹³C of up to ?3.5 ‰. In these areas, δ¹³C below ?12 ‰ suggests a 100 % C₃ grass and/or leafy plant (browse) diet while animals from higher latitude have lower δ¹³C. Animals from semi-arid areas have δ¹⁸O of 34–40 ‰, while grazers from temperate areas have lower values (~28–30 ‰). Three patterns with implications for palaeoenvironmental reconstruction emerge: (1) all species in semi-arid areas regularly browse to supplement limited grass resources; (2) all species within an environmental zone have similar carbon and oxygen isotope compositions, meaning data from different kangaroo species can be pooled for palaeoenvironmental investigations; (3) relatively small regional environmental differences can be distinguished when δ¹³C and δ¹⁸O data are used together. These data demonstrate that diet–isotope and climate–isotope relationships should be evaluated in modern ecosystems before application to the regional fossil record.     

Intraspecific variation in photosynthetic responses of trebouxioid lichens with reference to the activity of a carbon-concentrating mechanism

The photosynthetic responses of a range of trebouxioid lichens were investigated to determine whether variations in net assimilation rates shown by populations of the same species collected from different habitats could be correlated with adjustments in carbon-concentrating mechanism (CCM) activity. The activity of a CCM was inferred from the high affinity for CO₂ [i.e. low CO₂ compensation point (Γ); low external CO₂ concentration at which half-maximal assimilation rates are reached (K _{0.5 CO2})], the release of a pool of accumulated dissolved inorganic carbon (C_i) during light/dark transient measurements of CO₂ exchange and values for carbon isotope discrimination intermediate between those characteristic of C₃ and C₄ terrestrial plants. Higher net and gross assimilation rates were expressed by lichens collected from shaded woodland habitats. The higher rates were not accounted for by variations in chlorophyll content. Lichens with high assimilation rates also showed an increased affinity for CO₂ as demonstrated by low CO₂ compensation points and K _0.5 values and the magnitude of the C_i pool accumulated upon illumination and released after darkening of the thalli. However, there was no correlation between assimilation rates and organic matter or instantaneous carbon isotope discrimination measurements, with the latter remaining roughly consistent whatever the provenance or species of the lichen material. The data are discussed with reference to significant environmental factors which are likely to control photosynthesis in the habitats studied. Received: 5 April 1997 / Accepted: 9 September 1997