Geographical distribution and chorology of grasses in the Arabian Peninsula

Arid regions of Saudi Arabia occupy most of the area of the Arabian Peninsula. These areas are at the meeting position of plants from Mediterranean, Irano-Turanian, Saharo-Arabian, and Sudanian phytogegraphical regions. Geomorphology of the area reveals a wide diversity of landforms including coastal lines, desert plains, and high mountains. Grasses are well represented in the flora of Saudi Arabia and form an appropriate group for studying the relation of grass distribution, chorology, and photosynthetic pathways. In this paper, geographical distribution of C₃ and C₄ grasses was studied in an area extending between latitude 17°N and latitude 31°N. Two regions were recognized in the study area, namely; a (relatively) cold region north of latitude 24°N with ample winter rainfall, and a hot region south of latitude 24°N with scarce summer rainfall. Work involved field observations and collection of grass species in the study area. Work also depended on published carbon discrimination values of grasses and biochemical analysis of C₄ species subtypes. Climatic conditions in the study area vary considerably, and the distribution of grass species was found to follow patterns that reveal adaptive advantages of different photosynthetic pathways. Grass species in the cold northern region with ample winter rainfall are generally C₃ grasses belonging mainly to Mediterranean/Irano-Turanean chorotypes. C₃ grass species found in the southern hot region were recorded at high altitudes of southern mountains characterized by low temperatures. Grass species recorded at low altitudes in the south hot region with scarce summer rainfall were mainly C₄ grasses belonging to Tropical and Saharo-Arabian-Sudanean chorotypes. Pronounced spatial variations of temperature profoundly control the geographical distribution of C₃ and C₄ grasses. Low temperatures in the northern cold region and at high altitudes of the southern hot region limit the occurrence of C₄ grasses and shift the ecological balance in favor of C₃ grasses. Results are discussed in terms of heat sensitivity of the CO₂ carboxylating enzyme of C₃ grasses and high temperature optima for CO₂ assimilation of C₄ grasses. Results are also discussed in comparison with geographical distribution of grasses in other parts of the world.     

Photosynthetic pathways,distribution, and ecological characteristics of grass species in egypt

Summary This study is based both on our own results, especially δ¹³C analyses, and on data in the literature. Of the 225 species of Poaceae reported from Egypt, 105 species show the C₃ and 120 the C₄ type of photosynthetic CO₂ fixation. Winter annual and perennial grasses active in winter are mainly C₃ species, while summer annuals and the other perennials are mainly C₄ species. The percentage of C₃ species decreases with decreasing latitude. This seems to be mainly related to increasing temperatures. The C₃ grass species are found preferentially in the Mediterranean, Irano-Turanian, Mediterranean/Irano-Turanian, and Saharo-Arabian chorotypes, while the C₄ species are mainly found in the Sudanian, Saharo-Arabian/Sudanian and Tropical chorotypes. In Egypt, the NADP⁺-ME and NAD⁺-ME subtypes of C₄ photosynthesis are found in about equal numbers of C₄ species, while the PCK subtype is relatively rare.     

Relative nutritional quality of C3 and C4 grasses for a graminivorous lepidopteran,Paratrytone melane (Hesperiidae)

Summary We tested the hypothesis that C₄ grasses are inferior to C₃ grasses as host plants for herbivorous insects by measuring the relative performance of larvae of a graminivorous lepidopteran, Paratrytone melane (Hesperiidae), fed C₃ and C₄ grasses. Relative growth rates and final weights were higher in larvae fed a C₃ grass in Experiment I. However, in two additional experiments, relative growth rates and final weights were not significantly different in larvae fed C₃ and C₄ grasses. We examined two factors which are believed to cause C₄ grasses to be of lower nutritional value than C₃ grasses: foliar nutrient levels and nutrient digestibility. In general, foliar nutrient levels were higher in C₃ grasses. In Experiment I, protein and soluble carbohydrates were digested from a C₃ and a C₄ grass with equivalent efficiencies. Therefore, differences in larval performance are best explained by higher nutrient levels in the C₃ grass in this experiment. In Experiment II, soluble carbohydrates were digested with similar efficiencies from C₃ and C₄ grasses but protein was digested with greater efficiency from the C₃ grasses. We conclude (1) that the bundle sheath anatomy of C₄ grasses is not a barrier to soluble carbohydrate digestion and does not have a nutritionally significant effect on protein digestion and (2) that P. melane may consume C₄ grasses at compensatory rates.     

Differences in drought sensitivities and photosynthetic limitations between co-occurring C3 and C4 (NADP-ME) Panicoid grasses

Background and Aims

The success of C₄ plants lies in their ability to attain greater efficiencies of light, water and nitrogen use under high temperature, providing an advantage in arid, hot environments. However, C₄ grasses are not necessarily less sensitive to drought than C₃ grasses and are proposed to respond with greater metabolic limitations, while the C₃ response is predominantly stomatal. The aims of this study were to compare the drought and recovery responses of co-occurring C₃ and C₄ NADP-ME grasses from the subfamily Panicoideae and to determine stomatal and metabolic contributions to the observed response.

Methods

Six species of locally co-occurring grasses, C₃ species Alloteropsis semialata subsp. eckloniana, Panicum aequinerve and Panicum ecklonii, and C₄ (NADP-ME) species Heteropogon contortus, Themeda triandra and Tristachya leucothrix, were established in pots then subjected to a controlled drought followed by re-watering. Water potentials, leaf gas exchange and the response of photosynthetic rate to internal CO₂ concentrations were determined on selected occasions during the drought and re-watering treatments and compared between species and photosynthetic types.

Key Results

Leaves of C₄ species of grasses maintained their photosynthetic advantage until water deficits became severe, but lost their water-use advantage even under conditions of mild drought. Declining C₄ photosynthesis with water deficit was mainly a consequence of metabolic limitations to CO₂ assimilation, whereas, in the C₃ species, stomatal limitations had a prevailing role in the drought-induced decrease in photosynthesis. The drought-sensitive metabolism of the C₄ plants could explain the observed slower recovery of photosynthesis on re-watering, in comparison with C₃ plants which recovered a greater proportion of photosynthesis through increased stomatal conductance.

Conclusions

Within the Panicoid grasses, C₄ (NADP-ME) species are metabolically more sensitive to drought than C₃ species and recover more slowly from drought.     

Le vie fotosintetiche C3, C4 e CAM nel contesto ecologico

Abstract

Ecological aspects of C₃, C₄ and CAM photosynthetic pathways. - Three different photosynthetic CO₂ fixation pathways are known to occur in higher plants. However all three pathways ultimately depend on the Calvin-Benson cycle for carbon reduction. The oxygenase activity of RuBP carboxilase is responsible for photorespiratory CO₂ release. Both C₄ and CAM pathways behave as a CO₂ concentrating mechanism which prevent photorespiration. The CO₂-concentrating mechanism in C₄ plants is based on intracellular symplastic transport of C₄ dicarboxylic acids from mesophyll-cells to the adjacent bundle-sheath cells. On the contrary in CAM plants the CO₂-concentrating mechanism is based on the intracellular transport of malic acid into and out of the vacuole.

The C₄ photosynthetic pathway as compared to the C₃ pathway permits higher rates of CO₂ fixation in high light and high temperature environments at low costs in terms of water loss, given the stability of the photosynthetic apparatus under such conditions.

CAM is interpreted as an adaptation to arid environments because it enables carbon assimilation to take place at very low water costs during the night when the evaporative demand is low. Nevertheless many aquatic species of Isoetes and some relatives are CAM, suggesting the adaptive role of CAM to environments which become depleted in CO₂.

The photosynthetic carbon fixation pathway certainly contributes to the ecological success of plants in different environments. However the distribution of plants may also reflect their biological history. On the other hand plants with different photosynthetic pathways coexist in many communities and tend to share resources in time. In any case some generalizations are possible: C₄ plants enjoy an ecological advantage in hot, moist, high light regions while the majority of species in desert environments are C₃; CAM plants are more frequent in semiarid regions with seasonal rainfall, coastal fog deserts, and in epiphytic habitats in tropical rain forests.     

Quo vadis C4? An ecophysiological perspective on global change and the future of C4 plants

C₄ plants are directly affected by all major global change parameters, often in a manner that is distinct from that of C₃ plants. Rising CO₂ generally stimulates C₃ photosynthesis more than C₄, but C₄ species still exhibit positive responses, particularly at elevated temperature and arid conditions where they are currently common. Acclimation of photosynthesis to high CO₂ occurs in both C₃ and C₄ plants, most notably in nutrient-limited situations. High CO₂ aggravates nitrogen limitations and in doing so may favor C₄ species, which have greater photosynthetic nitrogen use efficiency. C₄ photosynthesis is favored by high temperature, but global warming will not necessarily favor C₄ over C₃ plants because the timing of warming could be more critical than the warming itself. C₃ species will likely be favored where harsh winter climates are moderated, particularly where hot summers also become drier and less favorable to C₄ plant growth. Eutrophication of soils by nitrogen deposition generally favors C₃ species by offsetting the superior nitrogen use efficiency of C₄ species; this should allow C₃ species to expand at the expense of C₄ plants. Land-use change and biotic invasions are also important global change factors that affect the future of C₄ plants. Human exploitation of forested landscapes favors C₄ species at low latitude by removing woody competitors and opening gaps in which C₄ grasses can establish. Invasive C₄ grasses are causing widespread forest loss in Asia, the Americas and Oceania by accelerating fire cycles and reducing soil nutrient status. Once established, weedy C₄ grasses can prevent woodland establishment, and thus arrest ecological succession. In sum, in the future, certain C₄ plants will prosper at the expense of C₃ species, and should be able to adjust to the changes the future brings. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of irradiance and temperature on photosynthesis in C3, C4 and C3/C4 Panicum species

Species in the Laxa and Grandia groups of the genus Panicum are adapted to low, wet areas of tropical and subtropical America. Panicum milioides is a species with C₃ photosynthesis and low apparent photorespiration and has been classified as a C₃/C₄ intermediate. Other species in the Laxa group are C₃ with normal photorespiration. Panicum prionitis is a C₄ species in the Grandia group. Since P. milioides has some leaf characteristics intermediate to C₃ and C₄ species, its photosynthetic response to irradiance and temperature was compared to the closely related C₃ species, P. laxum and P. boliviense and to P. prionitis. The response of apparent photosynthesis to irradiance and temperature was similar to that of P. laxum and P. boliviense, with saturation at a photosynthetic photo flux density of about 1 mmol m^-2 s^-1 at 30°C and temperature optimum near 30°C. In contrast, P. prionitis showed no light saturation up to 2 mmol m^-2 s^-1 and an optimum temperature near 40°C. P. milioides exhibited low CO₂ loss into CO₂-free air in the light and this loss was nearly insensitive to temperature. Loss of CO₂ in the light in the C₃ species, P. laxum and P. boliviense, was several-fold higher than in P. milioides and increased 2- to 5-fold with increases in temperature from 10 to 40°C. The level of dark respiration and its response to temperature were similar in all four Panicum species examined. It is concluded that the low apparent photorespiration in P. milioides does not influence its response of apparent photosynthesis to irradiance and temperature in comparison to closely related C₃ Panicum species.Abbreviations AP apparent photosynthesis - I CO₂ compensation point - gl leaf conductance; gm, mesophyll conductance - PPFD photosynthetic photon flux density - PR apparent photorespiration rate - RuBPC sibulose bisphosphate carboxylase     

Occurrence of the suberized lamella in leaves of grasses of different photosynthetic types. I. In parenchymatous bundle sheaths and PCR (“Kranz”) sheaths

Summary Leaf blades of 42 grasses (Poaceae) have been examined ultrastructurally for the occurrence of a suberized lamella in walls of parenchymatous bundle sheaths and PCR (= Kranz) sheaths in both large and small vascular bundles. The sample includes species from a range of major grass taxa, and represents all photosynthetic types found in the grasses. Three grasses with unusual C₄ leaf anatomy were also included:Alloteropsis semialata, Aristida biglandulosa, Arundinella nepalensis. The presence of a suberized lamella in PCR cell walls was perfectly correlated with photosynthetic type. All PEP-carboxykinase type and NADP-malic enzyme type C₄ species examined possessed a suberized lamella in outer tangential and radial walls, but with variable presence in inner tangential walls. PCR cells of bothAlloteropsis semialata andArundinella nepalensis also possessed a suberized lamella. A lamella was totally absent from parenchymatous bundle sheath cells of the C₃ species examined (5 spp.) and ofPanicum milioides, a C₃-C₄ intermediate. It was also absent from PCR cells of NAD-malic enzyme type C₄ species (14 spp.) andAristida biglandulosa. The results are discussed in relation to the leakage of CO₂ from PCR cells, and to differences between C₄ types in ¹³C values, chloroplast position in PCR cells, and other anatomical characteristics.     

Water-use efficiency of one C3 and two C4 grasses in response to varying soil moisture and herbage-removal levels in a seasonally dry tropical region

In a seasonally dry tropical region the water use efficiency (WUE) of three grasses (C₃ winter annualPolypogon monspeliensis, C₄ perennialDichanthium annulatum and C₄ warm seasonal annualEchinochloa colonum) was evaluated during summer and winter under nine experimental conditions (3 soil moisture×3 herbage removal). Generally leaf water status and transpiration rate decreased with soil moisture stress and increased with clipping intensity. During winter the transpiration rate of Dichanthium was much lower than that of Polypogon and its own rate in summer. Both soil moisture stress and clipping intensity increased the WUE in all instances. Despite differences in photosynthetic type, growing season and life form, these grasses exhibited broadly similar positive relationships, across nine treatments for WUE: soil moisture stress, and water consumption: production. The range of WUE (g. mm^–1) calculated on TNP through the nine treatments was: summer—Dichanthium 2.9–10.0, Echinochloa 2.0–6.7; winter—Dichanthium 4.3–36.3, Polypogon 1.9–12.0.     

Nitrogen distribution and leaf area indices in relation to photosynthetic nitrogen use efficiency in savanna grasses

Leaf photosynthetic characteristics, distribution patterns of nitrogen content per unit leaf area (n_L) and leaf area production per unit n_Lwere measured in natural stands of a C₄ grass (Hyparrhenia rufa) from the seasonal savannas and of a C₄grass (Paspalum fasciculatum) and two C₃grasses (Leersia hexandra and Hymenachne amplexicaulis) from the flooded savannas in central Venezuela. Daily rates of canopy photosynthesis (P_cD) as well as the optimal leaf area production per unit n_Lat which P_cDfor a given total amount of nitrogen in the canopy (i.e., canopy-PNUE) is maximized were also calculated. The C₃and C₄species from the flooded savannas had similar light saturated rates of photosynthesis per unit n_L(i.e. leaf-PNUE) and similar canopy-PNUEs which was in strong contrast with previous studies. Especially H. rufa but also L. hexandra and H. amplexicaulis had leaf- and canopy-PNUEs which were considerably higher than the values calculated for most other species with the same photosynthetic pathway (i.e., C₃or C₄). In contrast to previous studies, differences in the light gradient in the canopy between stands only partially explained differences in N distribution. Measured leaf area indices were greater and the average n_L values were consequently smaller than the calculated optima. There was, however, a very strong linear correlation between the optimal and actual average n_Lindicating that even though the model overestimated average n_L, it did predict the differences in leaf area production per unit nitrogen – the inverse average n_L– very well. This result strongly indicates that leaf area production per unit of leaf nitrogen increases with leaf-PNUE and decreases with the extinction coefficient for light. Grass species from seasonal savannas have extremely high leaf-PNUEs and thus optimally produce large amounts of leaf area per unit n_L. This helps explain how stands of these species may have high leaf area indices and achieve high photosynthetic productivity despite the very low nutrient availability at which they grow.     

Photosynthesis pathways,ecological characteristics,and the geographical distribution of the Cyperaceae in Japan

Summary The nature of the photosynthetic pathways of Cyperaceae found in Japan were investigated on the basis of Kranz anatomy, the CO₂ compensation concentration and previously reported data. Among 301 species (96% of all cyperaceous species recorded in the region), 58 species were classified as being C₄ plants. These C₄ species were scattered among the tribes Fimbristylideae, Lipocarpheae, Cypereae and Rhynchosporeae in the subfamily Cyperoideae. The genera Cyperus, Eleocharis and Rhynchospora included, in Japan, both C₃ and C₄ species within a single genus. Using these data, an analysis was made of the ecological characteristics and geographical distribution of the C₃ and C₄ species in Japan. Although cyperaceous species grow in markedly different environments, the majority were found in wet and aquatic areas (61%) or shaded areas, such as forest floors (20%). Most of the C₃ species were also hygrophytes (58%) and forest-living species (25%), and C₃ species growing in mesic and dry areas were relatively rare. The C₄ species inhabited wet and aquatic (75%), mesic (13%) and dry areas (6%) and showed marked ecological characteristics with respect to soil-moisture conditions, unlike other C₄ plants, although they were absent from shaded habitats. In order to determine the climatic factors that influence the relative floristic abundance of C₃ and C₄ members of the Cyperaceae in Japan, the ratios of number of C₄ species to the total number of members of Cyperaceae (C₄ percentage) in 16 representative locales were examined in terms of various climatic variables. There were strong positive correlations between the C₄ percentage and temperature. Among the C₃ groups of three subfamilies, there were different distributional trends for various temperature regimes. The C₃ subfamily Caricoideae increased its relative contribution to the cyperaceous flora with a decrease in mean annual temperature, while the C₃ subfamily Sclerioideae exhibited the opposite pattern. The C₃ group of the subfamily Cyperoideae did not show any marked change in pattern along temperature gradients, unlike the two other C₃ subfamilies, and seemed to be heterogeneous in terms of its response to temperature. The relationships between the C₄ biochemical subtypes and ecological characteristics are also discussed.     

Photosynthetic pathway,chilling tolerance and cell sap osmotic potential values of grasses along an altitudinal gradient in Papua New Guinea

Summary A total of 22 grass species were examined from 5 sites spanning the altitudinal range 1550–4350 m.a.s.l. The presence of the C₃ or C₄ photosynthetic pathway was determined from ¹³C values and chilling tolerance was assessed on the basis of electrolyte leakage from leaf slices incubated on melting ice. Most of the grasses studied at the lower altitude sites of 1550 m.a.s.l. (annual mean of daily minimum temperature, 14.6° C) and 2600 m.a.s.l. (9.4° C) possessed C₄ photosynthesis and were chill-sensitive. The single except ion was Agrostis avenacea, a montane chill-resistant C₃ species which occurred at 2600 m.a.s.l. The three species apparently most sensitive to chilling were Ischaemum polystachyum, Paspalum conjugatum and Saccharum robustum, all occurring at 1550 m.a.s.l. At the higher altitude sites of 3280 (5.6° C), 3580 (4.0° C) and 4350 (–0.7°C) m.a.s.l., most of the grasses exhibited C₃ photosynthesis and were chill-resistant. However, an Upland population of the C₄ species, Miscanthus floridulus was found at 3280 m.a.s.l. which had acquired chill-resistance as confirmed by additional in vivo variable chlorophyll fluorescence measurements. Cell sap osmotic potential values of the upland grasses at altitudes of 3280–4350 m.a.s.l. were lower (–8.1 to –19.8 bars) than values in grasses from 1550 and 2600 m.a.s.l. (–3.9 to –7.5 bars) due mainly to the presence of non-electrolyte osmoticants, which may be involved in frost avoidance mechanism(s).Abbreviations ABA abscisic acid - F_R the maximal rate of rise of induced chlorophyll fluorescence - s osmotic potential     

Performance of a generalist grasshopper on a C<Subscript>3</Subscript> and a C<Subscript>4</Subscript> grass: compensation for the effects of elevated CO<Subscript>2</Subscript> on plant nutritional quality

The increasing CO₂ concentration in Earths atmosphere is expected to cause a greater decline in the nutritional quality of C₃ than C₄ plants. As a compensatory response, herbivorous insects may increase their feeding disproportionately on C₃ plants. These hypotheses were tested by growing the grasses Lolium multiflorum C₃) and Bouteloua curtipendula C₄) at ambient (370 ppm) and elevated (740 ppm) CO₂ levels in open top chambers in the field, and comparing the growth and digestive efficiencies of the generalist grasshopper Melanoplus sanguinipes on each of the four plant × CO₂ treatment combinations. As expected, the nutritional quality of the C₃ grass declined to a greater extent than did that of the C₄ grass at elevated CO₂; protein levels declined in the C₃ grass, while levels of carbohydrates (sugar, fructan and starch) increased. However, M. sanguinipes did not significantly increase its consumption rate to compensate for the lower nutritional quality of the C₃ grass grown under elevated CO₂. Instead, these grasshoppers appear to use post-ingestive mechanisms to maintain their growth rates on the C₃ grass under elevated CO₂. Consumption rates of the C₃ and C₄ grasses were also similar, demonstrating a lack of compensatory feeding on the C₄ grass. We also examined the relative efficiencies of nutrient utilization from a C₃ and C₄ grass by M. sanguinipes to test the basis for the C₄ plant avoidance hypothesis. Contrary to this hypothesis, neither protein nor sugar was digested with a lower efficiency from the C₄ grass than from the C₃ grass. A novel finding of this study is that fructan, a potentially large carbohydrate source in C₃ grasses, is utilized by grasshoppers. Based on the higher nutrient levels in the C₃ grass and the better growth performance of M. sanguinipes on this grass at both CO₂ levels, we conclude that C₃ grasses are likely to remain better host plants than C₄ grasses in future CO₂ conditions.     

Stable isotope characteristics across narrow savanna/woodland ecotones in Wolfe Creek Meteorite Crater,Western Australia

The stable isotopic composition (δ¹³C) of sediments from lakes are frequently analyzed to reconstruct the proportion of the regional vegetation that used either the C₃ or C₄ photosynthetic pathways, often without conducting a detailed survey of the current local vegetation. We performed a study on the modern vegetation composition within the Wolfe Creek Meteorite Crater to complement our future paleoecological investigation of the crater. A bull’s-eye pattern exists where C₄ grasses dominate an outer ring and salt tolerant species, including shrubs, herbs, chenopods, and halophytic algae, dominate the inner pan of the crater. The ecotone between the inner and outer zones is narrow and occupied by tall (>7 m) Acacia ampliceps, with some C₄ grasses in the understory. Along with the highest water table and most saline soils the center of the crater has C₃ plants present with the highest δ¹³C and δ¹⁵N values. The range of δ¹³C and δ¹⁵N values from the analysis of surface soil organic matter (OM) was much smaller compared with the range of values from plant materials implying that either: (1) the current plant OM has not yet been integrated into the soils, or (2) processes within the soil have acted to homogenize isotopic variability within the crater. The application of a two end member mixing model to calculate %C₄ and %C₃ biomass from the δ¹³C of surface soil OM was complicated by: (1) the crater containing both a dry habitat with C₄ grasses and a central pan with C₄ halophytic plants and, (2) the large variation in the δ¹³C of the plants and soil OM.     

Implications of interveinal distance for quantum yield in C<Subscript>4</Subscript> grasses: a modeling and meta-analysis

The distance between veins has the potential to affect photosynthesis in C₄ grasses because photon capture and photosynthetic carbon reduction are primarily restricted to vascular bundle sheath cells (BSC). For example, BSC density should increase as interveinal distance (IVD) decreases, and thus IVD may influence photon capture and photosynthesis in C₄ grasses. The objective of this study is to evaluate the potential importance of IVD to the function of C₄ grasses, and a literature survey is conducted to test the hypothesis that quantum yield of photosynthesis () increases with decreasing IVD. First, a meta-analysis of and IVD values obtained for 12 C₄ grass species supports this hypothesis as and IVD are significantly negatively correlated (r=–0.61). Second, a regression of carbon isotope discrimination () versus IVD was conducted and the regression equation was used in a simple biochemical model that relates to and leakage of CO₂ from the BSC. The modeling analysis also supports the hypothesis that decreases with increasing IVD in C₄ grasses. C₄ grasses are virtually absent from shaded habitats, and the biochemical model is employed to examine the implications of IVD for shade-tolerance in C₄ grasses. The model predicts that only those species with uncommonly small IVD values are able to tolerate prolonged shade.     

Photosynthetic Pathways,Life Forms,and Reproductive Types for Forage Species Along the Desertification Gradient on Hunshandake Desert,North China

Floristic compositions, life forms, reproductive types for forage species, and their responses to desertification in Hunshandake desert were studied. 164 species, in 30 families and 94 genera, were identified with C₃ (137 species), C₄ (25 species), and CAM (2 species) photosynthesis. Of the 25 C₄ species, 76 % were grasses and Chenopodiaceae species (hereafter chenopods). This suggests that the C₄ species mainly occurred in a few families in the desert region. The reduction of C₃ species and the increase of C₄ species with desertification indicated that C₄ species might have higher tolerance to environmental stresses (e.g. dry and poor soil). Relatively more hemicrytophyte and therophyte forms in the desert are related to the local temperate climate and vegetation dynamics. Relatively greater proportions of C₄/C₃ and clonal species/sexual species at mobile dune showed that the C₄ species and clonal species could make greater contribution to sand land restoration in the Hunshandake desert.     

Leaf gas exchange characteristics and water- and nitrogen-use efficiencies of dominant grass and tree species in a West African savanna

Whereas leaf gas exchange properties are important to assess carbon and water fluxes in ecosystems worldwide, information of this type is scarce for savanna species. In this study, gas exchange characteristics of 2 C₄ grass species (Andropogon canaliculatus and Hyparrhenia diplandra) and 2 C₃ tree species (Crossopteryx febrifuga and Cussonia arborea) from the West-African savanna of Lamto (Ivory Coast) were investigated in the field. Measurements were done in order to provide data to allow the parameterization of biochemically-based models of photosynthesis (for C₄ and C₃ plant metabolic types) and stomatal conductance ; and to compare gas exchange characteristics of coexisting species. No systematic difference was found between grass and tree species for reference stomatal conductance, under standard environmental conditions, or stomatal response to incident light or vapour pressure deficit at leaf surface. Conversely, grass species displayed higher water (1.5-2 fold) and nitrogen (2-5 fold) photosynthetic use efficiencies (WUE and NUE, ratio of net photosynthesis to transpiration and leaf nitrogen, respectively). These contrasts were attributed to the CO₂ concentrating mechanism of C₄ plants. When looking within plant life forms, no important difference was found between grass species. However, significant contrasts were found between tree species, Cussonia showing higher NUE and reference stomatal conductance than Crossopteryx. These results stress the need to account for functional diversity when estimating ecosystem carbon and water fluxes. In particular, our results suggest that the tree/grass ratio, and also the composition of the tree layer, could strongly affect WUE and NUE at the ecosystem scale in West African savannas.     

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.     

Photosynthetic carbon metabolism of the cool-temperate C4 grass Spartina anglica Hubb.

The aim of this work was to describe the photosynthetic carbon metabolism of the cooltemperate C₄ grass Spartina anglica. With the exception of pyruvate, phosphate dikinase and pyruvate kinase, the maximum catalytic activities in leaves of putative enzymes of the C₄ cycle of a phosphoenolpyruvate-carboxykinase C₄ plant were considerably in excess of the observed, steady-state rate of photosynthesis, and were comparable with the maximum catalytic activities of key enzymes of the reductive pentose-phosphate pathway. Radioactive carbon from ¹⁴CO₂ supplied to attached leaves during steady-state photosynthesis appeared first in malate and aspartate from which it moved to intermediates of the reductive pentose-phosphate pathway, and then to sucrose. These experiments show that photosynthetic carbon metabolism in this cool-temperate C₄ plant is similar to that of C₄ plants of hotter climates.