Photosynthetic Pathway Types of Forage Species Along Grazing Gradient from the Songnen Grassland,Northeastern China

Photosynthetic pathway types (C₃ and C₄ species) and their dynamics along grazing gradient were determined for 42 plant species in 30 genera and 13 families from the Songnen grassland, Northeastern China. Of the total, 10 species in 9 genera and 4 families had C₄ photosynthesis; 32 species in 21 genera and 12 families had C₃ photosynthesis. The proportion of C₄ species in total plants and C₄/C₃ increased with grazing intensity, and peaked in overgrazed plot. Most of the increased C₄ species (6 of 10) along the grazed gradient were annual grasses and halophytes. This indicated that the C₄ species had greater capacity to tolerate environmental stresses (e.g. drought and saline) caused by animal grazing in the Songnen grassland, Northeastern China.     

Photosynthetic Pathways and Life Form Types for Native Plant Species from Hulunbeier Rangelands,Inner Mongolia,North China

Photosynthetic pathway Types (C₃, C₄, and CAM) and life forms of native species from Hulunbeier rangelands, north China were studied. Of the total 258 species, 216 species in 132 genera and 42 families had C₃ photosynthetic pathway, including dominant herbs, e.g. Stipa baicalensis Roshev. and Leymus chinensis (Trin.) Tzvel., Filifolium sibiricum Kitam. and Arudinella hirta (Thunb.) Koidz. 38 species in 28 genera and 10 families were found with C₄ photosynthesis, and 4 species in 2 genera and 1 family had CAM photosynthetic pathway. The occurrence of C₄ species was common in Gramineae and Chenopodiaceae, and the two families were leading ones within C₄ plants. More than 52 % of the total 258 species were in H form, 21 % in Th form, 19 % in G form; the other life form Types, e.g. Ch, M, N, and HH, formed less than 3 %. 68 % of C₄ species were in Th form and 24 % in H form, indicating that these Types were the dominant life forms for C₄ species in the rangeland region. The occurrence of C₄ species was closely related with plant habitats, disturbed lands had the highest C₄ abundance (55 % of the total C₄ species), followed by grasslands and sandy soil, and forests had the lowest C₄ abundance (8 %). Hence the occurrence of C₄ species could be efficient indicator for rangeland dynamics in Hulunbeier rangelands.     

The C4 Photosynthetic Pathway and Life Forms in Grassland Species from North China

C₄ photosynthetic pathway and life form were determined for 159 species in 71 genera and 13 families in the grassland of North China. 45 % of the C₄ species were found in Graminae, 19 % in each of Cyperaceae and Chenopodiaceae. More than 51 % of these C₄ species were in therophyta and 36 % hemicryptophyta, while fewer species were in nanophanerophyta (9 %) or geophyta (5 %). The numbers of C₄ species and their life forms were closely related with grassland deterioration and succession in North China. This indicated that the C₄ species had greater capacity to tolerate environmental stress (e.g. drought and salinity) caused by animal grazing and cultivation.     

C<Subscript>3</Subscript> and C<Subscript>4</Subscript> photosynthetic pathways and life form types for native species from agro-forestry region,Northeastern China

Of the total 570 species, 194 species in 116 genera and 52 families were found with C₃ photosynthesis, 24 species in 17 genera and 6 families with C₄ photosynthesis, and 2 species in 1 genera and 1 family with CAM photosynthesis. 90 % of the total species can be found in Changbai Mountain flora, more a half (69 %) in North China flora, and about 1/3 in Mongolian flora and Xinan flora, respectively. The occurrence of C₄ species was not as common as that in adjacent grasslands and deserts, but relatively more than in the adjacent forests. Of the total 24 C₄ species, 63 % C₄ species (15 of 24) was found in Gramineae. Nine life form types can be found, reflecting the moist climate in the region, especially the occurrence of epiphyte and liana forms. Relatively more geophyte life form plants suggested the winter in the region was much colder than in grasslands. These indicated that both ecological studies and land management decisions must take into account plant photosynthetic pathway and life form patterns, for both of them are closely related to climatic changes and land use.     

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 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.     

Photosynthetic Pathway Types in Rangeland Plant Species from Inner Mongolia,North China

Photosynthetic pathway types, based on ¹³C measurements, were determined for 125 species in 95 genera and 32 families growing in rangelands from Inner Mongolia. Of the total species, 4 species from 3 genera and 2 families had C₄ photosynthesis (2 species in Gramineae and 2 in Chenopodiaceae) and 118 species from 90 genera and 31 families had C₃ photosynthesis. The number of C₄ species differed significantly among four rangeland sites, 4 species in desert, 3 species in steppe, but no C₄ species were identified in meadow and dune. Six species [e.g. Agriophyllum arenarium Bieb., Bassia dasyphylla O. Kuntze, Saussurea japonica (Thunb.) DC.] earlier identified as C₄ species using the enzyme ratio method were found as C₃ species using the carbon isotope ratios (¹³C). Hence the enzyme ratio method for C₃ and C₄ identification may not always be reliable. The ¹³C values of 3 species of Crassulaceae, which had been considered as CAM species, differed remarkably [–25.79 for Sedum aizoon L., –24.42 for Osostachys fimbriatus (Turcz.) Berger, and –16.97 for O. malacophyllus (Pall.) Fisch], suggesting that the use of ¹³C method as a diagnosis for CAM photosynthetic pathway type may not always be reliable and supplementary measurements are needed.     

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.     

Photosynthetic and morphological functional types for native species from mixed prairie in Southern Saskatchewan, Canada

Photosynthetic pathways (e.g. C₃, C₄) and morphological functional types (e.g. trees, shrubs, high perennial grasses, perennial forbs) were identified for the native species from the Saskatchewan mixed prairie, using the data from references published between 1950 and 2003. Of the total 219 identified species in 145 genera and 45 families, 208 species in 137 genera and 44 families were found with C₃ photosynthesis, and most of these species are dominants (e.g. Agropyron dasystachyum Hook. and Stipa spartea var. curtiseta Hitchc.). 11 species in 10 genera and 3 families were identified with C₄ photosynthesis (e.g. Atriplex argentea Nutt., Andropogon scoparius Michx., Boutelou gracilis Lag., Calamovilfa longifolia Hook.). The amount of total identified C₄ species in the region is much less than that from the South Dakota mixed prairie (27 species). Gramineae is the leading family with C₄ photosynthesis (8 species), Chenopodiaceae ranks the second (2 species). Relatively less forb types [50 % perennial forbs (PEF) and 12 % annual forbs (ANF)] and more graminoid types (25 %) composition suggested that the rangelands in the region are relatively stable. Lacking of the knowledge on the optimal traits for PFTs classification in the region, further studies (e.g. C₃ and C₄ plant identification and optimal trait selection) are needed to explore the relationships between PFTs and vegetation variations, as well as land-use and climate changes.     

δ<Superscript>13</Superscript>C values,photosynthetic pathways,and plant functional types for some plants from saline meadows,Northeastern China

Based on stable carbon isotope ratio (δ¹³C) measurements, photosynthetic pathway types were determined for 61 species in 54 genera and 24 families of flowering plants from the saline meadows of Northeastern China. Of these total vascular plants, 18 species in 17 genera from 6 families were found to have C₄ photosynthesis; 43 species in 38 genera from 20 families had C₃ photosynthesis. Six dicotyledonous species exhibited C₄ pathway, 12 monocotyledonous species were found with C₄ photosynthesis. The dicotyledonous C₄ species had relative greater mean δ¹³C value and less total carbon content than both monocotyledonous C₄ and C₃ species. Most dicotyledonous C₄ species were annual forbs and halophytes. Some C₄ species had been previously documented, but their δ¹³C values varied remarkably from those of the present study. Even though there are some fluctuations for the δ¹³C values of some C₄ species, δ¹³C value was still more reliable for C₃ and C₄ identification than the use of the enzyme ratio method and of low CO₂ compensation concentration.     

基于结构方程模型分析森林草原带草本物种丰富度对景观因子的响应

受损景观是由不同比例的植被斑块组成的镶嵌体,阐明植被斑块的景观结构特征对物种多样性的影响,有助于提高受损景观物种多样性保护。在塞罕坝自然保护区选取38个天然植被斑块,其中包括12个草地、11个灌木林和15个天然次生林。根据光合作用的不同途径将草本物种划分为C_3和C_4功能群。选取斑块面积、形状指数、隔离度指数及每个斑块500 m缓冲区内的森林和草地比例作为景观因子。通过结构方程模型探讨C_3、C_4草本物种丰富度与景观因子的相互关系。斑块面积(2.18—74.06 hm~2)与C_3、C_4草本及总物种丰富度分布格局均具有显著的正相关关系(P0.05),且对C_3草本的影响最大;形状指数(1.06—3.11)、隔离度指数(33.51—327.65)对C_3、C_4草本及总物种丰富度影响不显著(P0.05);C_3、C_4草本及总物种丰富度与毗邻斑块草地比例(4.20%—64.95%)呈正相关,而与森林比例(35.05%—95.80%)呈负相关。研究区斑块面积和毗邻斑块植被构成是影响C_3、C_4草本植物的主要因素。在破碎化景观中保存面积大的天然植被斑块及提高毗邻斑块草地比例是保护关键C_3、C_4草本植物物种的有效途径。     

Geographic distributions and physiological characteristics of co-existing Flaveria species in south-central Mexico

The genus Flaveria consists of 23 species with significant variation in photosynthetic physiologies. We tested whether photosynthetic pathway variation in seven co-existing Flaveria species corresponds to geographic distributions or physiological performance in C₃, C₄, and intermediate species growing under natural conditions in south-central Mexico. We found that Flaveria pringlei (C₃) was the most widely distributed species with multiple growth habits. Numerous populations of Flaveria kochiana (C₄), a recently described species with a previously unknown distribution, were located in the Mixtec region of Oaxaca. Flaveria cronquistii (C₃) and Flaveria ramosissima (C₃-C₄) were only located in the Tehuacán Valley region while Flaveria trinervia (C₄) was widely distributed. Only one population of Flaveria angustifolia (C₃-C₄) and Flaveria vaginata (C₄-like) were located near Izúcar de Matamoros. Midday leaf water potential differed significantly between Flaveria species, but did not vary according to growth habit or photosynthetic pathway. The quantum yield of photosystem II did not vary between species, despite large differences in leaf nitrogen content, leaf shape, plant size and life histories. We did not find a direct relationship between increasing C₄ cycle characteristics and physiological performance in the Flaveria populations examined. Furthermore, C₃ species were not found at higher elevation than C₄ species as expected. Our observations indicate that life history traits and disturbance regime may be the primary controllers of Flaveria distributions in south-central Mexico.     

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.     

Photosynthetic pathway and morphological functional types in the vegetation from North-Beijing agro-pastoral ecotone, China

Photosynthetic pathways (C₃, C₄, and CAM) and morphological functional types were identified for the species from vegetation in agro-pastoral ecotone, North Beijing. 792 vascular plant species (nearly half of the total species in the ecotone), in 66 families and 317 genera, were identified with C₃, C₄, and CAM photosynthesis (Table 1). 710 species (90 % of the identified species in Table 1) in 268 genera and 64 families were found with C₃ photosynthesis, 68 species (9 % of the total identified species) in 40 genera and 7 families with C₄ photosynthesis, and 14 species in 4 genera and 1 family with CAM photosynthesis. Gramineae is the leading family with C₄ photosynthesis (43 species), Cyperaceae ranks the second (16 species) followed by Chenopodiaceae (5 species). The significant increase of C₄ proportion (C₄/total species) with land deterioration suggested the plants of this type are remarkably responsive to land use in the ecotone. 792 species were classified into nine morphological functional types and the changes of most of these types (e.g. perennial forbs (PEF), annual grasses (ANG), and annual forbs (ANF)) were consistent with habitats and vegetation dynamics in the agro-pastoral ecotone. Hence the photosynthetic pathways, combined with the morphological functional types, are efficient indications for studying the linkage between species and ecosystems in the ecotone.     

Occurrence and altitudinal pattern of C<Subscript>4</Subscript> plants on Qinghai Plateau,Qinghai province,China

The natural occurrence and altitudinal pattern of species with C₄ photosynthesis were investigated on Qinghai Plateau, Qinghai province by using stable carbon isotopes in plant leaves and using additional data from references. A total of 58 species belonging to 10 families and 34 genera were identified using C₄ photosynthetic pathway, which is only 1.66 % of total 3 500 plant species in Qinghai province. The leading two families, i.e. Gramineae (23 species) and Chenopodiaceae (22 species) contain 77.6 % of all C₄ plants in the studied area. The number of C₄ species increased from 1 600 to 2 400 m a.s.l. and then decreases quickly till 4 400 m a.s.l. with one half of C₄ species distributing from 2 200 to 2 800 m a.s.l. (48 %). Eight plant species were found above 4 000 m a.s.l., but the distribution of these species is limited to the south of Qinghai province (low latitude area) where annual mean temperature is above 0 °C, suggesting that low temperature may generally limit the distribution of C₄ plants.     

Carbon isotope composition as a tool to control the quality of herbs and medicinal plants

Isotope screening is a simple test for determining the photosynthetic pathway used by plants. The scope of this work was to classify the photosynthetic type of some herbs and medicinal plants through studies of the carbon isotope composition (δ¹³C). Also, we propose the use of carbon isotope composition as a tool to control the quality of herbs and medicinal plants. For studies of δ¹³C, δ¹³C‰ = [R (sample)/R (standard) − 1] × 10⁻³, dry leaves powdered in cryogenic mill were analyzed in a mass spectrometer coupled with an elemental analyzer for determining the ratio R = ¹³CO₂/¹²CO₂. In investigation of δ¹³C of 55 species, 23 botanical families, and 44 species possessed a C₃ photosynthetic type. Six species found among the botanical families Euphorbiaceae and Poaceae were C₄ plants, and 5 species found among the botanical families Agavaceae, Euphorbiaceae, and Liliaceae possessed CAM-type photosynthesis. Carbon isotope composition of plants can be used as quality control of herbs and medicinal plants, allowing the identification of frauds or contaminations. Also, the information about the photosynthetic type found for these plants can help in introducing and cultivating exotic and wild herbs and medicinal plants.     

A comparative immunocytochemical localization study of phosphoenolpyruvate carboxylase in leaves of higher plants

The intracellular localization of phosphoenolpyruvate (PEP) carboxylase in plants belonging to the C₄, Crassulacean acid metabolism (CAM) and C₃ types was invetigated using an immunocytochemical method with an immune serum raised against the sorghum leaf enzyme. The plants studied were sorghum, maize (C₄ type), kalanchoe (CAM type), french bean, and spinach (C₃ type). In the green leaves of C₄ plants, it was shown that the carboxylase was located in the mesophyll and stomatic cells, being largely cytosolic in the mesophyll cells. Similarly, in CAM plants, the enzyme was found mainly outside the chloroplasts. In contrast, in C₃ plants, the PEP carboxylase appeared to be distributed between the cytosol and the chloroplasts of foliar parenchyma. Examination of sections from etiolated leaves showed fluorescence emission from etioplasts and cytosol for the parenchyma of french bean as well as for the bundle sheath and mesophyll of sorghum leaves. This data indicated that during the greening process photoregulation and evolution of PEP carboxylase is dependent on the tissue and on the metabolic type of the plant considered.Abbreviations CAM Crassulacean acid metabolism - PEP phosphoenolpyruvate     

Leaf carbon isotope ratios of plants from a subtropical monsoon forest

Summary Carbon isotope ratios were used to survey the distribution of photosynthetic pathways among taxa, the relationship between photosynthetic pathway and habitat light levels, and the relationship between intercellular CO₂ levels of C₃ plants and habitat light levels within a subtropical monsoon forest in southern China. Of 128 species, most (94) possessed the C₃ photosynthetic pathway; 33 species possessed the C₄ pathway and all of these were restricted to high light locations. There was one epiphytic CAM species. The C₃ species were classified as occurring in open, intermediate, and closed canopy sites. Among C₃ species, carbon isotope ratios tended to become more negative with decreasing light availability in the habitat.C.I.W.D.P.B. Pub no 931     

Ecological selection pressures for C4 photosynthesis in the grasses

Grasses using the C₄ photosynthetic pathway dominate grasslands and savannahs of warm regions, and account for half of the species in this ecologically and economically important plant family. The C₄ pathway increases the potential for high rates of photosynthesis, particularly at high irradiance, and raises water-use efficiency compared with the C₃ type. It is therefore classically viewed as an adaptation to open, arid conditions. Here, we test this adaptive hypothesis using the comparative method, analysing habitat data for 117 genera of grasses, representing 15 C₄ lineages. The evidence from our three complementary analyses is consistent with the hypothesis that evolutionary selection for C₄ photosynthesis requires open environments, but we find an equal likelihood of C₄ evolutionary origins in mesic, arid and saline habitats. However, once the pathway has arisen, evolutionary transitions into arid habitats occur at higher rates in C₄ than C₃ clades. Extant C₄ genera therefore occupy a wider range of drier habitats than their C₃ counterparts because the C₄ pathway represents a pre-adaptation to arid conditions. Our analyses warn against evolutionary inferences based solely upon the high occurrence of extant C₄ species in dry habitats, and provide a novel interpretation of this classic ecological association.