Climate-driven changes in shoot density and shoot biomass in Leymus chinensis (Poaceae) on the North-east China Transect (NECT)

Aim Climate‐driven changes affecting ecosystem primary production have been well documented for many vegetation types, while the effects of climate on plant populations remains unclear. Herein, we address the relationships between climatic variables and shoot density, reproductive allocation and shoot biomass in Leymus chinensis on a large‐scale climatic gradient in 2000. Location Nine sites experiencing similar light regimes, but differing in longitude, precipitation and altitude were selected on the North‐east China Transect (NECT) from 115° to 124°E, around a latitude of 43.5°N. Methods Densities of total, vegetative and reproductive shoots and of shoot biomass were measured twice over the growing season in each site. Climatic data were taken from the climate database of the Laboratory of Quantitative Vegetation Ecology, Institute of Botany, Chinese Academy of Sciences and from the local weather stations throughout the NECT. Results Densities of total, vegetative and reproductive shoots increased significantly from the west to the east and from dry to moist along the NECT, and were strongly correlated with annual precipitation (r² = 0.934, 0.943 and 0.863, respectively) and an aridity index (r² = 0.809, 0.816 and 0.744, respectively). The average total shoot density at the east end (470/m²) was about three times that at the west (160/m²). Reproductive allocation and shoot biomass for both vegetative and reproductive shoots increased with precipitation and declined with the aridity index along the NECT. There were positive correlations between shoot biomass and annual precipitation for vegetative shoots (P < 0.05, R² = 0.604) and March precipitation for reproductive shoots (P < 0.05, R² = 0.533), respectively. Main conclusions These findings suggest that L. chinensis adjusts to decreasing precipitation/increasing aridity by alterations in shoot density, reproductive allocation and shoot biomass along the drought gradient of the NECT.     

The occurrence of C4 plants and their morphological functional types in the vegetation of Xinjiang, China

Floristic composition, morphological functional types and habitat distributions for C₄ species were studied in Xinjiang, North-western China. 89 species, in 9 families and 41 genera, were identified with C₄ photosynthesis. 48 % of these C₄ species were found in Monocotyledoneae, e.g. Cyperaceae (5 species), Gramineae (38 species), the other 52 % was in Dicotyledoneae, e.g. Chenopodiaceae (29 species), Amaranthaceae (7 species), and Polygonaceae (5 species). Compared with those in semi-arid grasslands in North China and tropical forests in South China, more plant families were found with the occurrence of C₄ plants in this arid region. Relatively higher annual species (63 %), shrubs (18 %), and Chenopodiaceae C₄ species (33 %) compositions were the primary characteristics for the C₄ species occurring in Xinjiang, and this was remarkably related with its arid environment. More Chenopodiaceae C₄ species occurring in the region suggested that this type of C₄ species may have higher capacity to fit the air and soil droughts. There was a strong relationship between C₄ occurrence and habitat distributions, more than half of the total 89 C₄ species were found in disturbed and cultivated lands and early stages of vegetation successions, indicating C₄ occurrence was not only related with climate changes, but also with land uses and vegetation dynamics.     

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.     

C4 plants in the vegetation of Mongolia: their natural occurrence and geographical distribution in relation to climate

The natural geographical occurrence, carbon assimilation, and structural and biochemical diversity of species with C₄ photosynthesis in the vegetation of Mongolia was studied. The Mongolian flora was screened for C₄ plants by using ¹³C/¹²C isotope fractionation, determining the early products of ¹⁴CO₂ fixation, microscopy of leaf mesophyll cell anatomy, and from reported literature data. Eighty C₄ species were found among eight families: Amaranthaceae, Chenopodiaceae, Euphorbiaceae, Molluginaceae, Poaceae, Polygonaceae, Portulacaceae and Zygophyllaceae. Most of the C4 species were in three families: Chenopodiceae (41 species), Poaceae (25 species) and Polygonaceae, genus Calligonum (6 species). Some new C₄ species in Chenopodiaceae, Poaceae and Polygonaceae were detected. C₄ Chenopodiaceae species make up 45% of the total chenopods and are very important ecologically in saline areas and in cold arid deserts. C₄ grasses make up about 10% of the total Poaceae species and these species naturally concentrate in steppe zones. Naturalized grasses with Kranz anatomy,of genera such as Setaria, Echinochloa, Eragrostis, Panicum and Chloris, were found in almost all the botanical-geographical regions of Mongolia, where they commonly occur in annually disturbed areas and desert oases. We analyzed the relationships between the occurrence of C₄ plants in 16 natural botanical-geographical regions of Mongolia and their major climatic influences. The proportion of C₄ species increases with decreasing geographical latitude and along the north-to-south temperature gradient; however grasses and chenopods differ in their responses to climate. The abundance of Chenopodiaceae species was closely correlated with aridity, but the distribution of the C₄ grasses was more dependent on temperature. Also, we found a unique distribution of different C₄ Chenopodiaceae structural and biochemical subtypes along the aridity gradient. NADP-malic enzyme (NADP-ME) tree-like species with a salsoloid type of Kranz anatomy, such as Haloxylon ammodendron and Iljinia regelii, plus shrubby Salsola and Anabasis species, were the plants most resistant to ecological stress and conditions in highly arid Gobian deserts with less than 100 mm of annual precipitation. Most of the annual C₄ chenopod species were halophytes, succulent, and occurred in saline and arid environments in steppe and desert regions. The relative abundance of C₃ succulent chenopod species also increased along the aridity gradient. Native C₄ grasses were mainly annual and perennial species from the Cynodonteae tribe with NAD-ME and PEP-carboxykinase (PEP-CK) photosynthetic types. They occurred across much of Mongolia, but were most common in steppe zones where they are often dominant in grazing ecosystems. Received: 17 March 1999 / Accepted: 1 November 1999     

Response of photosynthesis of different plant functional types to environmental changes along Northeast China Transect

Net photosynthesis (P_n), transpiration (E), stomatal conductance (g_s), internal CO₂ concentration (C_i), and water use efficiency (WUE) were examined on 215 species from eight plant functional types (PFTs) along a precipitation gradient in northeast China (the Northeast China Transect, or NECT). Among the eight PFTs, meadow steppe grasses had the highest rates of net photosynthesis and forest grasses the lowest and the following order of P_n was noted: meadow steppe grasses >typical steppe grasses >steppe shrubs >desert grasses >forest trees >forest shrubs >desert shrubs >forest grasses (P<0.05). Transpiration tended to be the highest in the steppe grasses and lowest in forest shrubs. Transpiration also decreased rapidly with the appearance of C₃ desert species at the desert end. The forest tree PFT had lower P_n, E, g_s than the steppe PFTs, whereas WUE values were somewhat greater in the forest tree PFT than the desert shrubs and grasses. Low C_i values along the steppe section (from 400 to 1100 km, east to west) indicated the presence of C₄ species. Of all the PFTs, only shrubs and herbs were noted at all points along the transect. No clear relationship between P_n, E, g_s, WUE of herb and shrub PFTs and annual precipitation was noted – low values were found at both the high and low precipitation ends of the transect. Highest values were noted when precipitation was intermediate. Received: 28 October 1998 / Accepted: 10 May 1999     

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.     

The occurrence of C4 photosynthesis in Yunnan province, a tropical region in South-western China

Floristic composition, morphological functional types, and altitudinal distribution pattern for C₄ species were studied in Yunnan province, South-western China. 159 species, in 6 families and 60 genera, were identified with C₄ photosynthesis. 93 % of these C₄ species were found in Monocotyledoneae, e.g. Cyperaceae (18 species), Gramineae (129 species), and Commelinaceae (1 species), the other 7 % was in Dicotyledoneae, e.g. Amaranthaceae (5 species), Portulacaceae (4 species), and Chenopodiaceae (2 species). Hence C₄ plants mainly occurred in very few families in the tropical region. Compared with those in semi-arid grasslands and arid deserts in North China, more C₄ grasses and much less Chenopodiaceae C₄ species occurred in the tropical region. This indicates the physiological responses of C₄ plants from the two families are very different. Chenopodiaceae C₄ species may be more fit semi-arid and arid environments, while C₄ grasses are more fit the moist tropical conditions. There was a strong relationship between C₄ distribution and altitude in the tropical region. Altitudinal distribution pattern for C₄ species in the region was consistent with altitude, climate, and habitats.     

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.     

European plants with C4 photosynthesis: geographical and taxonomic distribution and relations to climate parameters

A survey of C₄ plants in Europe was performed with 216 species based on information in the literature and new studies. C₄ species were found in 10 families: the eudicots Amaranthaceae, Chenopodiaceae, Euphorbiaceae, Molluginaceae, Nyctaginaceae, Polygonaceae, Portulacaceae and Zygophyllaceae and the monocots Cyperaceae and Poaceae. The majority of the C₄ species belong to four families, Amaranthaceae (23), Chenopodiaceae (65), Cyperaceae (27) and Poaceae (88). In central and southern Europe, the abundance of native C₄ plants varied between 44 and 88% of total C₄ plants occurring, the rest being invasive C₄ species. The occurrence of total C₄ species, C₄ monocots and C₄ Chenopodiaceae was assessed for five major phyto‐geographical regions of Europe (north‐west, north‐east, central, south‐west, and south‐east). The abundance of C₄ plants of total C₄ dicots, C₄ Chenopodiaceae, total C₄ monocots, C₄ Poaceae and C₄ Cyperaceae was related to the climatic variables of annual mean daily temperature, annual precipitation and DeMartonne's aridity index. The abundance of total C₄ plants decreases with increasing temperature and expression of aridity (decreasing aridity index) and is not correlated with precipitation. The abundance of total C₄ dicots and C₄ Chenopodiaceae is correlated with precipitation and aridity but not temperature, whereas the abundance of total C₄ monocots, C₄ Poaceae and C₄ Cyperaceae is correlated with temperature and aridity but not precipitation. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 283–304.     

Photosynthetic Pathways and Life Forms in Different Grassland Types from North China

Photosynthetic pathways (C₄, C₃, and CAM species) and plant life forms of three grassland types in North China were compared. Of the total 201 species, 144 species in 78 genera and 34 families had C₃ photosynthetic pathway, 56 species in 35 genera and 11 families had C₄ photosynthetic pathway, and 1 species had CAM photosynthetic pathway. The number of C₄ species in Songnen meadow was 70–80 % greater than that in Xilinguole steppe and Hunshandak desert grassland, but that for C₃ species did not differ significantly among the three grassland types. The number of therophytes in the Songnen meadow was relatively greater than that of the other two grassland types, but that of hemicryptophytes was lower. Thus the distribution of C₄ species and plant life form is probably related to precipitation.     

C<Subscript>4</Subscript> plants in the deserts of China: occurrence of C<Subscript>4</Subscript> photosynthesis and its morphological functional types

C₄ photosynthetic pathway and morphological functional types were determined for 104 species in 45 genera and 10 families from the deserts of China. 67 C₄ species (64.4 %) were found in Dicotyledoneae (e.g. Chenopodiaceae, Polygonaceae, and Amaranthaceae), the other 37 species were in Monocotyledoneae (e.g. Gramineae, Cyperaceae, and Commelinaceae). 36.5 % of the Chenopodiaceae species (predominantly members of the genera Anabasis, Atriplex, Kochia, Salsola, and Suaeda) identified in the desert regions were found with C₄ photosynthesis, which was about 48 % of the total C₄ species. Many C₄ species (58.7 %) were annuals (e.g. Amaranthus, Atriplex, Digitaria, Eragrostis, Kochia, and Salsola) and experienced long-term droughts, high temperature, and high irradiance. Relatively more shrub C₄ species (28 species of 104) were found in Chenopodiaceae (e.g. Anabasis, Camphorosma, Haloxylon, and Salsola) and Polygonaceae (e.g. Calligonum) in the desert regions. Most of shrub C₄ species with small leaf area were no more than 1 m in height and distributed in sandy soils. Composition of relatively more annual species, shrubs, and Chenopodiaceae C₄ species was the primary characteristic for the C₄ species occurrence in deserts, and this was remarkably related with the arid environmental conditions.     

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.     

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.     

Photosynthetic pathway variation among C4 grasses along a precipitation gradient in Argentina

Aim Based on the biochemical and physiological attributes of C₄ grasses, and on the close association between decarboxylation pathways and the taxa in which they evolved, the hypotheses tested were: (1) that C₄ grasses would become progressively more abundant as precipitation decreased, with grasses of the NADP‐me subtype more abundant in wetter sites and those of the NAD‐me subtype more common in arid regions; and (2) that the distribution of grass subfamilies would also be correlated with annual precipitation. Location The study was conducted along a precipitation gradient in central Argentina, from the eastern Pampas (>1000 mm year^?1) to the western deserts and semi‐deserts near the Andes (<100 mm year^?1). Methods Percentage of species and relative cover of C₃ and C₄ grasses (including C₄ subtypes) in local floras from 15 lowland sites of central Argentina were obtained from our own unpublished data and from recently published floristic surveys. Pearson correlation coefficients were obtained between grass distribution parameters and the available climatic data. Results The percentage of C₄ grasses increased towards the arid extreme and showed a strong negative correlation with annual rainfall (r = ?0.74, P < 0.01). Within the C₄ subtypes, the NADP‐me species showed a higher proportional representation at the wetter extreme, whereas the representation of NAD‐me species increased towards the more arid extreme. The relationship of PEP‐ck species with climatic parameters in central Argentina was less evident. The distributions of the Panicoideae and Chloridoideae subfamilies along the precipitation gradient were diametrically opposed, with the Panicoideae positively (r = 0.86, P < 0.001) and the Chloridoideae negatively (r = ?0.87, P < 0.001) correlated with annual precipitation. Main conclusions Our data are consistent with the broad observation that C₄ grasses tend to dominate in areas where the wet season falls in the warmer summer months. In agreement with previously reported results for Africa, Asia, Australia and North America, we describe here for the first time a significant relationship between annual precipitation and the prevalence of the NADP‐me and NAD‐me photosynthetic pathways along climatic gradients for the Neotropics. We also report for the first time that correlations between C₄ species and annual rainfall are stronger when the relative cover of grass species is considered. The association of grass subfamilies Panicoideae and Chloridoideae with rainfall is as strong as that recorded for the NADP‐me and NAD‐me variants, respectively, suggesting that characteristics other than decarboxylation type may be responsible for the geographic patterns described in this study.     

δ 13C variation of C3 and C4 plants across an Asian monsoon rainfall gradient in arid northwestern China

We have investigated carbon isotopic compositions of four plant genus/species, Bothriochloa ischaemum (C₄), Stipa bungeana (C₃), Lespedeza sp. (C₃) and Heteropappus less (C₃), along a precipitation gradient in northwest China in order to assess the impact of water availability on the carbon isotopic discrimination against ¹³C during carbon assimilation in this area. This information is necessary for reconstruction of paleovegetation, particularly paleo‐C₃/C₄ plant ratios using δ¹³C value of organic matter in loess and paleosols in the Chinese Loess Plateau. The δ¹³C of C₃ plants, as a group, exhibits a negative correlation with the annual precipitation amount with a total change and sensitivity of 5‰ and ?1.1‰/100 mm, respectively, for the precipitation range from 200 to 700 mm. The C₄ grass, B. ischaemum responds to aridity by decreasing 1.7‰ for over the precipitation range from 350 to 700 mm; the plant δ¹³C is significantly correlated with annual precipitation with a slope ?0.61‰/100 mm. This result implies that without considering the effect of water availability on the plant δ¹³C values, reconstruction of percent C₄ vegetation during the last glaciation can be overestimated by about a factor of two.     

C4 Plants in the Vegetation of Tibet,China: Their Natural Occurrence and Altitude Distribution Pattern

Floristic composition, life forms for C₄ species, and the pattern of altitude distribution were studied on Tibetan Plateau. 79 species, in 7 families and 46 genera, were identified with C₄ photosynthesis. 95 % of these C₄ species belong to Gramineae (51 species), Cyperaceae (14 species), and Chenopodiaceae (10 species), indicating that C₄ plants mainly occur in very few families (7 of 204) on the Tibetan Plateau. High altitude distribution for all the Chenopodiaceae C₄ species (> 3 000 m above sea level) suggests that plants of this kind have large tolerance to cold, dryness, and strong ultraviolet radiation. Most Gramineae and Cyperaceae C₄ species occurrences are consistent with extensive distribution of steppes and meadows in the vast flat of the central Plateau (1 000–3 000 m a.s.l.). Relatively high amount of hemicryptophyte form plants (44 %) in the region indicates that the vegetation, especially grassland, meadows, and steppe, are in good condition. There is a strong relationship between numbers of C₄ species and altitude in the Tibetan Plateau. Occurrence of C₄ species is significantly less in both high and low altitude plateaux in Tibet. Altitude distribution pattern for C₄ species in the region is not only consistent with the altitude and climate, but also with the vegetation types in altitude gradient.     

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.     

Impact of rainfall manipulations and biotic controls on soil respiration in Mediterranean and desert ecosystems along an aridity gradient

Spatially heterogeneous ecosystems form a majority of land types in the vast drylands of the globe. To evaluate climate‐change effects on CO₂ fluxes in such ecosystems, it is critical to understand the relative responses of each ecosystem component (microsite). We investigated soil respiration (R_s) at four sites along an aridity gradient (90–780 mm mean annual precipitation, MAP) during almost 2 years. In addition, R_s was measured in rainfall manipulations plots at the two central sites where ～30% droughting and ～30% water supplementation treatments were used over 5 years. Annual R_s was higher by 23% under shrub canopies compared with herbaceous gaps between shrubs, but R_s at both microsites responded similarly to rainfall reduction. Decreasing precipitation and soil water content along the aridity gradient and across rainfall manipulations resulted in a progressive decline in R_s at both microsites, i.e. the drier the conditions, the larger was the effect of reduction in water availability on R_s. Annual R_s on the ecosystem scale decreased at a slope of 256/MAP g C m^?2 yr^?1 mm^?1 (r²=0.97). The reduction in R_s amounted to 77% along the aridity gradient and to 16% across rainfall manipulations. Soil organic carbon (SOC) decreased with declining precipitation, and variation in SOC stocks explained 77% of the variation in annual R_s across sites, rainfall manipulations and microsites. This study shows that rainfall manipulations over several years are a useful tool for experimentally predicting climate‐change effects on CO₂ fluxes for time scales (such as approximated by aridity gradients) that are beyond common research periods. Rainfall reduction decreases rates of R_s not only by lowering biological activity, but also by drastically reducing shrub cover. We postulate that future climate change in heterogeneous ecosystems, such as Mediterranean and deserts shrublands will have a major impact on R_s by feedbacks through changes in vegetation structure.     

Co-function of C3-and C4-photosynthetic pathways in C3, C4 and C3-C4 intermediate Flaveria species

The potential for C₄ photosynthesis was investigated in five C₃-C₄ intermediate species, one C₃ species, and one C₄ species in the genus Flaveria, using ¹⁴CO₂ pulse-¹²CO₂ chase techniques and quantum-yield measurements. All five intermediate species were capable of incorporating ¹⁴CO₂ into the C₄ acids malate and aspartate, following an 8-s pulse. The proportion of ¹⁴C label in these C₄ products ranged from 50–55% to 20–26% in the C₃-C₄ intermediates F. floridana Johnston and F. linearis Lag. respectively. All of the intermediate species incorporated as much, or more, ¹⁴CO₂ into aspartate as into malate. Generally, about 5–15% of the initial label in these species appeared as other organic acids. There was variation in the capacity for C₄ photosynthesis among the intermediate species based on the apparent rate of conversion of ¹⁴C label from the C₄ cycle to the C₃ cycle. In intermediate species such as F. pubescens Rydb., F. ramosissima Klatt., and F. floridana we observed a substantial decrease in label of C₄-cycle products and an increase in percentage label in C₃-cycle products during chase periods with ¹²CO₂, although the rate of change was slower than in the C₄ species, F. palmeri. In these C₃-C₄ intermediates both sucrose and fumarate were predominant products after a 20-min chase period. In the C₃-C₄ intermediates, F. anomala Robinson and f. linearis we observed no significant decrease in the label of C₄-cycle products during a 3-min chase period and a slow turnover during a 20-min chase, indicating a lower level of functional integration between the C₄ and C₃ cycles in these species, relative to the other intermediates. Although F. cronquistii Powell was previously identified as a C₃ species, 7–18% of the initial label was in malate+aspartate. However, only 40–50% of this label was in the C-4 position, indicating C₄-acid formation as secondary products of photosynthesis in F. cronquistii. In 21% O₂, the absorbed quantum yields for CO₂ uptake (in mol CO₂·[mol quanta]^-1) averaged 0.053 in F. cronquistii (C₃), 0.051 in F. trinervia (Spreng.) Mohr (C₄), 0.052 in F. ramosissima (C₃-C₄), 0.051 in F. anomala (C₃-C₄), 0.050 in F. linearis (C₃-C₄), 0.046 in F. floridana (C₃-C₄), and 0.044 in F. pubescens (C₃-C₄). In 2% O₂ an enhancement of the quantum yield was observed in all of the C₃-C₄ intermediate species, ranging from 21% in F. ramosissima to 43% in F. pubescens. In all intermediates the quantum yields in 2% O₂ were intermediate in value to the C₃ and C₄ species, indicating a co-function of the C₃ and C₄ cycles in CO₂ assimilation. The low quantum-yield values for F. pubescens and F. floridana in 21% O₂ presumably reflect an ineffcient transfer of carbon from the C₄ to the C₃ cycle. The response of the quantum yield to four increasing O₂ concentrations (2–35%) showed lower levels of O₂ inhibition in the C₃-C₄ intermediate F. ramosissima, relative to the C₃ species. This indicates that the co-function of the C₃ and C₄ cycles in this intermediate species leads to an increased CO₂ concentration at the site of ribulose-1,5-bisphosphate carboxylase/oxygenase and a concomitant decrease in the competitive inhibition by O₂.Abbreviations PEP phosphoenolpyruvate - PGA 3-phosphoglycerate - RuBP ribulose-1,5-bisphosphate     

Morphological responses of Leymus chinensis (Poaceae) to the large-scale climatic gradient along the North-east China Transect (NECT)

A field experiment was conducted to examine the morphological variations of Leymus chinensis along the climatic gradient of the North‐east China Transect (NECT), from 115° to 125° E, in the vicinity of 43.5° N, in north‐eastern China. Ten sites selected for plant sampling along the gradient have approximately uniform theoretical radiation, but differ in precipitation and geographical elevation. The results of analyses showed that vegetative and reproductive shoot heights, flag leaf lengths and widths and seed numbers per inflorescence increased from the west to the east with precipitation, but decreased with aridity. Leaf lengths and widths for most leaf types from 115° to 124° E exhibited little variations. Significant correlations of plant heights, flag leaf lengths and widths and seed numbers per inflorescence with large‐scale climatic variables (e.g. annual precipitation, aridity) and geographical variation (longitude) found in this study indicated that climatic factors have significant effects on some morphological traits of L. chinensis along the NECT.