Artificial drainage and associated carbon fluxes (CO2/CH4) in a tundra ecosystem

Ecosystem flux measurements using the eddy covariance (EC) technique were undertaken in 4 subsequent years during summer for a total of 562 days in an arctic wet tundra ecosystem, located near Cherskii, Far-Eastern Federal District, Russia. Methane (CH₄) emissions were measured using permanent chambers. The experimental field is characterized by late thawing of permafrost soils in June and periodic spring floods. A stagnant water table below the grass canopy is fed by melting of the active layer of permafrost and by flood water. Following 3 years of EC measurements, the site was drained by building a 3 m wide drainage channel surrounding the EC tower to examine possible future effects of global change on the tundra tussock ecosystem. Cumulative summertime net carbon fluxes before experimental alteration were estimated to be about +15 g C m⁻² (i.e. an ecosystem C loss) and +8 g C m⁻² after draining the study site. When taking CH₄ as another important greenhouse gas into account and considering the global warming potential (GWP) of CH₄ vs. CO₂, the ecosystem had a positive GWP during all summers. However CH₄ emissions after drainage decreased significantly and therefore the carbon related greenhouse gas flux was much smaller than beforehand (475 ± 253 g C-CO₂-e m⁻² before drainage in 2003 vs. 23 ± 26 g C-CO₂-e m⁻² after drainage in 2005).     

植物排放N2O和CH4的研究

N₂O和CH₄是2种重要的温室气体, 但其排放源尚未得到充分鉴别。1990年和2006年先后报道植物能排放N₂O和CH₄, 并日益受到广泛的关注。然而, 迄今为止对植物排放这2种气体的研究均是分开单独进行的。该文以8种陆生草本植物为研究对象, 首次同步考察了新鲜离体植物地上部排放N₂O和CH₄的通量。研究结果表明: 8种植物均能排放这2种气体。其中, 黑麦草(Lolium perenne)、抱茎苦荬菜(Ixeridium sonchifolium)和菠菜(Spinacia oleracea)的CH₄通量较高, 分别为165.38、 52.28和21.64 ngCH₄·g^–1dw·h^–1; 抱茎苦荬菜、蒙古蒿(Artemisia mongolica)、大豆(Glycine max)和菠菜的N₂O通量较高, 分别为7.19、6.92、5.44和4.05 ngN₂O·g^–1dw·h^–1。研究结果不仅为植物本身既能排放N₂O又能排放CH₄在植物中可能具有普遍性提供了进一步的实验依据, 而且为深入研究其机理找到了几种适宜的植物种(如抱茎苦荬菜、菠菜)。     

C4 photosynthesis in a single C3 cell is theoretically inefficient but may ameliorate internal CO2 diffusion limitations of C3 leaves

Attempts are being made to introduce C₄ photosynthetic characteristics into C₃ crop plants by genetic manipulation. This research has focused on engineering single‐celled C₄‐type CO₂ concentrating mechanisms into C₃ plants such as rice. Herein the pros and cons of such approaches are discussed with a focus on CO₂ diffusion, utilizing a mathematical model of single‐cell C₄ photosynthesis. It is shown that a high bundle sheath resistance to CO₂ diffusion is an essential feature of energy‐efficient C₄ photosynthesis. The large chloroplast surface area appressed to the intercellular airspace in C₃ leaves generates low internal resistance to CO₂ diffusion, thereby limiting the energy efficiency of a single‐cell C₄ concentrating mechanism, which relies on concentrating CO₂ within chloroplasts of C₃ leaves. Nevertheless the model demonstrates that the drop in CO₂ partial pressure, pCO₂, that exists between intercellular airspace and chloroplasts in C₃ leaves at high photosynthetic rates, can be reversed under high irradiance when energy is not limiting. The model shows that this is particularly effective at lower intercellular pCO₂. Such a system may therefore be of benefit in water‐limited conditions when stomata are closed and low intercellular pCO₂ increases photorespiration.     

Mass spectrometric monitoring of gases (CO2, CH4, O2) in a mesotrophic peat core from Kopparås Mire, Sweden

Membrane inlet mass spectrometry was used to monitor dissolved gas concentrations (CO₂, CH₄ and O₂) in a mesotrophic peat core from Kopparås, Sweden. 1 A comparison of depth profiles (down to 22 cm) with an ombrotrophic peat core (Ellergower, SW Scotland) investigated previously, revealed major differences in gas concentrations. Thus methane reached concentrations more than twice as high (800 μM) at depths greater than 12 cm in the Kopparås core. As shown previously, the primary determinant of the depth of the oxic zone is the level of the water table. Whereas in the Scottish cores, mass spectrometric detectability of O₂ was confined to the first 3 cm below this level, in the Swedish core penetration of O₂ was greater (7 cm). CO₂ profiles were similar in cores from both locations. 2 A thick layer of Sphagnum mosses dominated the plant cover of the Swedish peat core. A poorly developed deep root system, as distinct from that of the vascular plant cover in Scottish cores, diminished gas exchange rates, and presumably aerobic methane oxidation at depth around roots. These characteristics may contribute to the development of discontinuities in gas profiles at depths greater 15 cm as upward gas transport is established predominantly by diffusion and/or ebullition in the Swedish core. 3 Monitoring gas concentrations at the peat surface and at 2 cm depth after changing water tables showed a delayed response of approximately 4 days as a result of the high water content and moisture‐regulating capacity of mosses. 4 Recovery processes at 2 cm depth after raising the water table revealed final production rates of dissolved CO₂ and CH₄ in the peat pore water between 0.8 and 4.4 μmol h^?1 L^?1 and between 0.1 and 1.7 μmol h^?1 L^?1, respectively. Higher production rates were found during the day, indicating a diurnal rhythm due to plant photosynthetic activity even at the low values of photosynthetically active radiation (PAR: 110 μmol s^?1 m^?2) used in the experimental set‐up. 5 In the water‐logged mesotrophic Kopparås core changes of dissolved gas concentrations (DGC) at 3 and 14 cm depth were surface temperature‐dependent rather than light dependent. This suggests that changes of air temperature alters the covering vegetation to increase the conductivity for dissolved gases through vascular plants and to facilitate gas transport by diffusion and/or ebullition.     

Effects of raised CO2 on potential CH4 production and oxidation in, and CH4 emission from, a boreal mire

1 In a glasshouse experiment we studied the effect of raised CO₂ concentration (720 p.p.m.) on CH₄ emission at natural boreal peat temperatures using intact cores of boreal peat with living vascular plants and Sphagnum mosses. After the end of the growing season half of the cores were kept unnaturally warm (17–20 °C). The potential for CH₄ production and oxidation was measured at the end of the emission experiment.
2 The vascular cores ('Sedge') consisted of a moss layer with sedges, and the moss cores (' Sphagnum ') of Sphagnum mosses (some sedge seedlings were removed by cutting). Methane efflux was 6–12 times higher from the Sedge cores than from the Sphagnum cores. The release of CH ₄ from Sedge cores increased with increasing temperature of the peat and decreased with decreasing temperature. Methane efflux from Sphagnum cores was quite stable independent of the peat temperatures.
3 In both Sedge and Sphagnum samples, CO₂ treatment doubled the potential CH₄ production but had no effect on the potential CH₄ oxidation. A raised concentration of CO₂ increased CH₄ efflux weakly and only at the highest peat temperatures (17–20 °C).
4 The results suggest that in cool regions, such as boreal wetlands, temperature would restrict decomposition of the extra substrates probably derived from enhanced primary production of mire vegetation under raised CO₂ concentrations, and would thus retard any consequent increase in CH₄ emission.     

Effects of N-fertilisation on CH4 oxidation and production, and consequences for CH4 emissions from microcosms and rice fields

The world's growing human population causes an increasing demand for food, of which rice is one of the most important sources. In rice production nitrogen is often a limiting factor. As a consequence increasing amounts of fertiliser will have to be applied to maximise yields. There is an ongoing discussion on the possible effects of fertilisation on CH₄ emissions. We therefore investigated the effects of N‐fertiliser (urea) on CH₄ emission, production and oxidation in rice microcosms and field experiments. In the microcosms, a substantial but short‐lived reduction of CH₄ emission was observed after N‐addition to 43‐d‐old rice plants. Methane oxidation increased by 45%, demonstrated with inhibitor measurements and model calculations based on stable carbon isotope data (δ¹³CH₄). A second fertilisation applied to 92‐d‐old plants had no effect on CH₄ emission rates. The positive effect of additional N on methanotrophic bacteria was also found in vitro for potential CH₄ oxidation rates in soil and root samples from the microcosm and field experiments, indicated by elevated initial oxidation rates and reduced lag‐phases. Fertilisation did not affect methane production in the microcosms. In the field, the effects were diverse: methane production was inhibited in the topsoil, but stimulated instead in the bulk soil. Stimulation occurred probably in the anaerobic food chain at the level of hydrolytic or fermenting bacteria, because acetate, a methanogenic precursor, increased simultaneously. Combining field, microcosm and laboratory experiments we conclude that any agricultural treatment improving the N‐supply to the rice plants will also be favourable for the CH₄ oxidising bacteria. However, N‐fertilisation had only a transient influence and was counter‐balanced in the field by an elevated CH₄ production. A negative effect of the fertilisation was a transient increase of N₂O emissions from the microcosms. However, integrating over the season the global warming potential (GWP) of N₂O emitted after fertilisation was still negligible compared to the GWP of emitted CH₄.     

Effects of free-air CO2 enrichment (FACE) on CH4 emission from a rice paddy field

Methane (CH₄) is a particularly potent greenhouse gas with a radiative forcing 23 times that of CO₂ on a per mass basis. Flooded rice paddies are a major source of CH₄ emissions to the Earth's atmosphere. A free‐air CO₂ enrichment (FACE) experiment was conducted to evaluate changes in crop productivity and the crop ecosystem under enriched CO₂ conditions during three rice growth seasons from 1998 to 2000 in a rice paddy at Shizukuishi, Iwate, Japan. To understand the influence of elevated atmospheric CO₂ concentrations on CH₄ emission, we measured methane flux from FACE rice fields and rice fields with ambient levels of CO₂ during the 1999 and 2000 growing seasons. Methane production and oxidation potentials of soil samples collected when the rice was at the tillering and flowering stages in 2000 were measured in the laboratory by the anaerobic incubation and alternative propylene substrates methods, respectively. The average tiller number and root dry biomass were clearly larger in the plots with elevated CO₂ during all rice growth stages. No difference in methane oxidation potential between FACE and ambient treatments was found, but the methane production potential of soils during the flowering stage was significantly greater under FACE than under ambient conditions. When free‐air CO₂ was enriched to 550 ppmv, the CH₄ emissions from the rice paddy field increased significantly, by 38% in 1999 and 51% in 2000. The increased CH₄ emissions were attributed to accelerated CH₄ production potential as a result of more root exudates and root autolysis products and to increased plant‐mediated CH₄ emissions because of the larger rice tiller numbers under FACE conditions.     

Models of carbon metabolism in C3-C4 intermediate plants as applied to the evolution of C4 photosynthesis

Abstract Models developed to explain the biphasic response of CO₂ compensation concentration to O₂ concentration and the C₃-like carbon isotope discrimination in C₃-C₄ intermediate species are used to characterize quantitatively the steps necessary in the evolution of C₄ photosynthesis. The evolutionary stages are indicated by model outputs, CO₂ compensation concentration and δ₁₃C value. The transition from intermediate plants to C₄ plants requires the complete formation of C₄ cycle capacity, expressed by the models as transition from C₄ cycle limitation by phosphoenolpyruvate (PEP) regeneration rate to limitation by PEP carboxylase activity. Other steps refer to CO₂ leakage from bundle sheath cells, to further augmentations of C₄ cycle components, to the repression of ribulose-1,5-bisphos-phate carboxylase in the mesophyll cells, and to a decrease in the CO₂ affinity of the enzyme. Possibilities of extending the suggested approach to other physiological characteristics, and the adaptive significance of the steps envisaged, are discussed.     

Fluxes of N2O and CH4 from soils of savannas and seasonally-dry ecosystems

Aim Savannas and seasonally‐dry ecosystems cover a significant part of the world's land surface. If undisturbed, these ecosystems might be expected to show a net uptake of methane (CH₄) and a limited emission of nitrous oxide (N₂O). Land management has the potential to change dramatically the characteristics and gas exchange of ecosystems. The present work investigates the contribution of warm climate seasonally‐dry ecosystems to the atmospheric concentration of nitrous oxide and methane, and analyses the impact of land‐use change on N₂O and CH₄ fluxes from the ecosystems in question. Location Flux data reviewed here were collected from the literature; they come from savannas and seasonally‐dry ecosystems in warm climatic regions, including South America, India, Australasia and Mediterranean areas. Methods Data on gas fluxes were collected from the literature. Two factors were considered as determinants of the variation in gas fluxes: land management and season. Land management was grouped into: (1) control, (2) ‘burned only’ and (3) managed ecosystems. The season was categorized as dry or wet. In order to avoid the possibility that the influence of soil properties on gas fluxes might confound any differences caused by land management, sites were grouped in homogeneous clusters on the basis of soil properties, using multivariate analyses. Inter‐ and intra‐cluster analysis of gas fluxes were performed, taking into account the effects of season, land management and main vegetation types. Results Soils were often acid and nutrient‐poor, with low water retention. N₂O emissions were generally very low (median flux 0.32 mg N₂O m^?2 day^?1), and no significant differences were observed between woodland savannas and managed savannas. The highest fluxes (up to 12.9 mg N₂O m^?2 day^?1) were those on relatively fertile soils with high air‐filled porosity and water retention. The effect of season on N₂O production was evident only when sites were separated in homogeneous groups on the basis of soil properties. CH₄ fluxes varied over a wide range (?22.9 to 3.15 mg CH₄ m^?2 day^?1, where the negative sign denotes removal of gas from the atmosphere), with an annual average daily flux of ?0.48 ± 0.96 (SD) mg CH₄ m^?2 day^?1 in undisturbed (control) sites. Land‐use change dramatically reduced this CH₄ sink. Managed sites were weak sinks of CH₄ in the dry season and became sources of CH₄ in the wet season. This was particularly evident for pastures. Burning alone did not reduce soil net CH₄ oxidation, but decreased N₂O production. Main conclusions Despite the low potential for N₂O production, both in natural and managed conditions, tropical seasonally‐dry ecosystems represent a significant source of N₂O (4.4 Tg N₂O year^?1) on a global scale, as a consequence of the large area they occupy. The same environments represent a potential CH₄ sink of 5.17 Tg CH₄ year^?1. However, assuming that c. 30% of the tropical land is converted to different uses, the sink would be reduced to 3.2 Tg CH₄ year^?1. The limited information on fluxes from Mediterranean ecosystems does not allow a meaningful scaling up.     

Methane emission from Texas rice paddy soils. 2. Seasonal contribution of rice biomass production to CH4 emission

Measurements focused on seasonal contribution of rice productivity to methane emission were made in three experiments conducted in Texas flooded paddy soils during 1994 and 1995 growing seasons. A total of five rice cultivars representing two distinct groups in methane emission were involved. Over a 10-week period after permanent flooding, total seasonal methane emission was positively correlated with rice above-ground biomass ( r ² = 0.845, n = 11). A very strong dependence of daily methane emission on above-ground vegetative biomass ( r ² = 0.887, n = 93) and on root biomass ( r ² = 0.816, n = 33) was also observed. Calculation from three developmental periods (vegetative, reproductive and ripening) of rice plant indicated that more than 75% of total seasonal methane was emitted during the last 5-week period in concert with reproductive and ripening stages, while rice biomass production during the same period amounted to ≈ 50% of the seasonal total. According to the correlation of cumulative methane emission with above-ground biomass increment between every two-week interval ( r ² = 0.490, n = 93, P = 0.000), the carbon released as methane is approximately equivalent to 3% and 4.5% of photosynthetically fixed carbon in the biomass for low and high emission cultivars, respectively. A further investigation showed that these fractions are related to plant growth and development. The carbon ratio of methane emitted to net photosynthetic production during vegetative, reproductive, and ripening periods averaged 0.9%, 3.6% and 7.9%, respectively, for low emission cultivars, and 2.0%, 5.0% and 8.3%, respectively, for high emission cultivars. Moreover, the ratio was strongly dependent on plant biomass, resulting in r 2 values from 0.775 to 0.907.     

The growth response of C4 plants to rising atmospheric CO2 partial pressure: a reassessment

Despite mounting evidence showing that C₄ plants can accumulate more biomass at elevated CO₂ partial pressure (p(CO₂)), the underlying mechanisms of this response are still largely unclear. In this paper, we review the current state of knowledge regarding the response of C₄ plants to elevated p(CO₂) and discuss the likely mechanisms. We identify two main routes through which elevated p(CO₂) can stimulate the growth of both well-watered and water-stressed C₄ plants. First, through enhanced leaf CO₂ assimilation rates due to increased intercellular p(CO₂). Second, through reduced stomatal conductance and subsequently leaf transpiration rates. Reduced transpiration rates can stimulate leaf CO₂ assimilation and growth rates by conserving soil water, improving shoot water relations and increasing leaf temperature. We argue that bundle sheath leakiness, direct CO₂ fixation in the bundle sheath or the presence of C₃-like photosynthesis in young C₄ leaves are unlikely explanations for the high CO₂-responsiveness of C₄ photosynthesis. The interactions between elevated p(CO₂), leaf temperature and shoot water relations on the growth and photosynthesis of C₄ plants are identified as key areas needing urgent research.     

Comparison of the specific activity of ribulose-1,5-bis-phosphate carboxylase-oxygenase from some C3 and C4 plants

The specific activity of ribulose-1,5-bisphosphate carboxylase-oxygenase (Rubisco, EC 4.1.1.39) was measured from the crude extracts of five C₃ plants consisting of wheat ( Triticum aestivum L. cv. Maris Mink), spinach ( Spinacia oleracea L.), pea ( Pisum sativum L. cv. Greenfeast), pumpkin ( Cucurbita pepo L. cv. Jättiläismeloni) and Ceratodon purpureus (Hedw.) Brid., and two C₄ plants, maize ( Zea mays L. ETA F₁) and sugar sorghum [ Sorghum saccharatum (L. emend, L.) Moench]. The amount of Rubisco in the crude extracts was estimated by polyacrylamide gel electro-phoresis with the Coomassie Brilliant Blue staining procedure. The amounts of the dye bound to the purified Rubisco of different higher plants were similar. The method gave a linear response for both purified enzyme and crude extracts, and the results agreed with those observed by immunochemical methods. The addition of positive effectors such as inorganic phosphate was necessary to obtain maximal activity in the crude extracts of all the studied plants except in that of maize. No significant differences in the specific carboxylase activity at 25°C were found between the C₃ and C₄ plants.     

Naturally low carbonic anhydrase activity in C4 and C3 plants limits discrimination against C18OO during photosynthesis

The ¹⁸O content of CO₂ is a powerful tracer of photosynthetic activity at the ecosystem and global scale. Due to oxygen exchange between CO₂ and ¹⁸O-enriched leaf water and retrodiffusion of most of this CO₂ back to the atmosphere, leaves effectively discriminate against ¹⁸O during photosynthesis. Discrimination against ¹⁸O ( Δ ¹⁸O) is expected to be lower in C₄ plants because of low c_i and hence low retrodiffusing CO₂ flux. C₄ plants also generally show lower levels of carbonic anhydrase (CA) activities than C₃ plants. Low CA may limit the extent of ¹⁸O exchange and further reduce Δ ¹⁸O. We investigated CO₂–H₂O isotopic equilibrium in plants with naturally low CA activity, including two C₄ (Zea mays, Sorghum bicolor) and one C₃ (Phragmites australis) species. The results confirmed experimentally the occurrence of low Δ ¹⁸O in C₄, as well as in some C₃, plants. Variations in CA activity and in the extent of CO₂–H₂O isotopic equilibrium ( θ _eq) estimated from on-line measurements of Δ ¹⁸O showed large range of 0–100% isotopic equilibrium ( θ _eq = 0–1). This was consistent with direct estimates based on assays of CA activity and measurements of CO₂ concentrations and residence times in the leaves. The results demonstrate the potential usefulness of Δ ¹⁸O as indicator of CA activity in vivo. Sensitivity tests indicated also that the impact of θ _eq < 1 (incomplete isotopic equilibrium) on ¹⁸O of atmospheric CO₂ can be similar for C₃ and C₄ plants and in both cases it increases with natural enrichment of ¹⁸O in leaf water.     

Characterization of purified leaf cytosolic pyruvate kinase from the C4 plant Cynodon dactylon

Cytosolic pyruvate kinase (EC 2.7.1.40) from leaves of the C₄ plant Cynodon dactylon (L.) Pers. was purified 56-fold to apparent homogeneity by polyethylene glycol fractionation and column chromatography including Q-Sepharose anion exchanger, ADP-Agarose and gel filtration. Nondenaturing PAGE of the final preparation resulted in a single protein band that co-migrated with the pyruvate kinase activity. Gel filtration and SDS-PAGE (± DTT) showed that this enzyme has a molecular mass of 200 kDa and is a homotetramer with a subunit molecular mass of 50 kDa. The subunits are not associated to each other with S-S bonds. The enzyme has a pH optimum of 6.2 and is heat stable. Typical Michaelis-Menten kinetics was obtained for both substrates, PEP and ADP, with K_m values of 64 and 235 μ M , respectively. Initial velocity studies indicated a sequential binding of the substrates to the enzyme.     

Ecosystem–atmosphere exchange of CH4 and N2O and ecosystem respiration in wetlands in the Sanjiang Plain, Northeastern China

Natural wetlands are critically important to global change because of their role in modulating atmospheric concentrations of CO₂, CH₄, and N₂O. One 4‐year continuous observation was conducted to examine the exchanges of CH₄ and N₂O between three wetland ecosystems and the atmosphere as well as the ecosystem respiration in the Sanjiang Plain in Northeastern China. From 2002 to 2005, the mean annual budgets of CH₄ and N₂O, and ecosystem respiration were 39.40 ± 6.99 g C m^?2 yr^?1, 0.124 ± 0.05 g N m^?2 yr^?1, and 513.55 ± 8.58 g C m^?2 yr^?1 for permanently inundated wetland; 4.36 ± 1.79 g C m^?2 yr^?1, 0.11 ± 0.12 g N m^?2 yr^?1, and 880.50 ± 71.72 g C m^?2 yr^?1 for seasonally inundated wetland; and 0.21 ± 0.1 g C m^?2 yr^?1, 0.28 ± 0.11 g N m^?2 yr^?1, and 1212.83 ± 191.98 g C m^?2 yr^?1 for shrub swamp. The substantial interannual variation of gas fluxes was due to the significant climatic variability which underscores the importance of long‐term continuous observations. The apparent seasonal pattern of gas emissions associated with a significant relationship of gas fluxes to air temperature implied the potential effect of global warming on greenhouse gas emissions from natural wetlands. The budgets of CH₄ and N₂O fluxes and ecosystem respiration were highly variable among three wetland types, which suggest the uncertainties in previous studies in which all kinds of natural wetlands were treated as one or two functional types. New classification of global natural wetlands in more detailed level is highly expected.     

Root exudation (net efflux of amino acids) may increase rhizodeposition under elevated CO2

Increases in atmospheric CO₂ concentration ([CO₂]) can lead to global climate change and theoretically could enhance carbon (C) deposition in soil, but data on this complex issue are contradictory. One approach for clarifying the diverse forces influencing plant‐derived C in the rhizosphere involves defining how elevated [CO₂] alters the fundamental process of C transfer from plant roots to the soil. We examine here how a step increase in [CO₂] affects the innate influx and efflux components of root exudation in axenic plants, as one foundation for understanding how climate change may affect rhizodeposition. Increasing [CO₂] from 425 to 850 μmol mol^?1 during short‐term trials enhanced shoot and root dry weight (P<0.01) of annual rye grass (Lolium multiflorum Lam.) and medic (Medicago truncatula L.) but had no effect on growth of maize (Zea mays L.). Root amino‐acid flux in the same plants changed only in maize, which increased the efflux rate (nmol g root fresh weight^?1 h^?1) of six amino acids (arginine, alanine, proline, tyrosine, lysine and leucine) significantly (P<0.05) under elevated [CO₂]. None of the three plant species altered the steady‐state concentration of 16 amino acids released into a hydroponic solution with changing [CO₂], apparently because amino‐acid influx rates, measured at 2.5 μm , consistently exceeded efflux rates. Indeed, plants recovered amino acids at rates 94–374% higher than they were lost from roots regardless of [CO₂]. These results indicate that, in theory, any effect of [CO₂] doubling on amino‐acid efflux can be offset by innately higher rates of influx. In practice, however, higher rates of amino‐acid cycling (i.e., efflux+influx) for each root segment (in C₄ maize) or from more root tissue (in the two C₃ species) should increase root exudation by plants exposed to elevated [CO₂] as additional amino acids would be adsorbed to soil particles or be taken up by soil microorganisms.     

Distribution of C3 and C4 grasses along an altitudinal gradient in Central Argentina

The distribution pattern of C₃ and C₄ grasses was studied in eight sites located between 350 m and 2100 m along an altitudinal gradient in Central Argentina. Of 139 taxa fifty-nine are C₃ and eighty C₄. Species of the C₃ tribes (Stipeae, Poeae, Meliceae, Aveneae, Bromeae and Triticeae) and C₃ Paniceae species increase in number at higher elevations; only one C₃ species was found below 650 m. C₄ Aristideae, Pappophoreae, Eragrostideae, Cynodonteae, Andropogoneae and Paniceae increase at lower altitudes. The floristic crossover point is at about 1500 m; the ground cover cross-over point is at about 1000 m. Analysis of the relationships between % C₄ species along the gradient and nine climatic and environmental variables showed the highest correlation with July mean temperature, but all temperature variables show highly significant correlations with % C₄. Correlation with annual rainfall is lower but also significant. These results are consistent with previous research showing the relative importance of C₄ grasses as temperature increases. C₃ species make a high contribution to relative grass coverage below the C₃/C₄ floristic crossover point but are rare below 1000 m.