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1.
Grasslands account for a large proportion of global terrestrial productivity and play a critical role in carbon and water cycling. Within grasslands, photosynthetic pathway is an important functional trait yielding different rates of productivity along environmental gradients. Recently, C3-C4 sorting along spatial environmental gradients has been reassessed by controlling for confounding traits in phylogenetically structured comparisons. C3 and C4 grasses should sort along temporal environmental gradients as well, resulting in differing phenologies and growing season lengths. Here we use 10 years of satellite data (NDVI) to examine the phenology and greenness (as a proxy for productivity) of C3 and C4 grass habitats, which reflect differences in both environment and plant physiology. We perform phylogenetically structured comparisons based on 3,595 digitized herbarium collections of 152 grass species across the Hawaiian Islands. Our results show that the clade identity of grasses captures differences in their habitats better than photosynthetic pathway. Growing season length (GSL) and associated productivity (GSP) were not significantly different when considering photosynthetic type alone, but were indeed different when considering photosynthetic type nested within clade. The relationship between GSL and GSP differed most strongly between C3 clade habitats, and not between C3-C4 habitats. Our results suggest that accounting for the interaction between phylogeny and photosynthetic pathway can help improve predictions of productivity, as commonly used C3-C4 classifications are very broad and appear to mask important diversity in grassland ecosystem functions.  相似文献   

2.
Grasses are ancestrally tropical understory species whose current dominance in warm open habitats is linked to the evolution of C4 photosynthesis. C4 grasses maintain high rates of photosynthesis in warm and water stressed environments, and the syndrome is considered to induce niche shifts into these habitats while adaptation to cold ones may be compromised. Global biogeographic analyses of C4 grasses have, however, concentrated on diversity patterns, while paying little attention to distributional limits. Using phylogenetic contrast analyses, we compared macro-climatic distribution limits among ~1300 grasses from the subfamily Panicoideae, which includes 4/5 of the known photosynthetic transitions in grasses. We explored whether evolution of C4 photosynthesis correlates with niche expansions, niche changes, or stasis at subfamily level and within the two tribes Paniceae and Paspaleae. We compared the climatic extremes of growing season temperatures, aridity, and mean temperatures of the coldest months. We found support for all the known biogeographic distribution patterns of C4 species, these patterns were, however, formed both by niche expansion and niche changes. The only ubiquitous response to a change in the photosynthetic pathway within Panicoideae was a niche expansion of the C4 species into regions with higher growing season temperatures, but without a withdrawal from the inherited climate niche. Other patterns varied among the tribes, as macro-climatic niche evolution in the American tribe Paspaleae differed from the pattern supported in the globally distributed tribe Paniceae and at family level.  相似文献   

3.
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.  相似文献   

4.
Wang  R.Z. 《Photosynthetica》2002,40(2):243-250
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.  相似文献   

5.
北京北部农牧交错区C4植物及其形态功能型和生境分析   总被引:1,自引:0,他引:1  
刘晓强  王仁忠 《生态学报》2006,26(5):1509-1515
根据野外调查和文献资料研究了北京北部农牧交错区C4植物的种类组成、形态功能型组成及其与生境的关系.该区共有野生C4植物68种,分布在7科,40个属,其中禾本科43种,莎草科16种,藜科5种.1年生(ANG和ANF)C4物种占C4植物总数的62%,它们在盐碱地、沙地、弃耕干扰地、湿地均超过这些生境C4植物分布的半数,在草地ANG和ANF超过30%;C4植物数量和C4/total在该地区的生境分布中有两个方向的变化:一是从草地到弃耕干扰地,两者均呈增加趋势,二是从草地到沙地和盐碱地呈现前者减少而后者增加的不同变化趋势,体现了农牧交错区植被退化的复杂性.  相似文献   

6.
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.  相似文献   

7.
Summary The implications of a reduced quantum yield (initial slope of the photosynthetic light response curve) in C4 plants and temperature dependence of quantum yield in C3 plants on total canopy primary production were investigated using computer simulations. Since reduced quantum yield represents the only known disadvantage of the C4 photosynthetic pathway, simulations were conducted with grass canopies (high LAI and hence photosynthesis in most leaves will be light-limited) to see if quantum yield is a significant factor in limiting the primary production and thus distributions of C4 grasses. Simulations were performed for three biogeographical or environmental conditions: the Great Plains region of North America, the Sonoran Desert of North America, and shade habitats. For all three cases, the simulations predicted either spatial or temporal gradients in the abundances of C4 grasses identical to the abundance patterns of C4 grasses observed in the field. It is thus concluded that while the C4 photosynthetic mechanism may be highly advantageous in specific environments, it may be disadvantageous in others.C.I.W.-D.P.B. Publication No. 598  相似文献   

8.
Measurements of the dependence of photosynthesis on light, CO2, and temperature are reported for two species of Encelia (Compositae) which differ in leaf pubescence and in geographical distribution. Encelia californica is glabrous and occurs in relatively mild, but arid habitats and Encelia farinosa is heavily pubescent and occurs in hot, arid habitats. Both species possess the C3 photosynthetic pathway. Under high irradiances and normal atmospheric conditions the two species have high photosynthetic rates, exceeding 3 nanomoles of CO2 per square centimeter per second (48 milligrams of CO2 per square decimeter per hour) and complete light saturation does not occur by full noon sunlight. The high photosynthetic capacity is related to a high efficiency of utilization of intercellular CO2 combined with high stomatal conductance. Leaf estimates of total soluble protein and fraction I protein are higher in these species than in most plants, although the proportion of fraction I protein is not higher. Both E. californica and E. farinosa attain a maximum rate of photosynthesis between 25 and 30 C, despite the fact that the two species grow in very different thermal habitats. Neither E. californica nor E. farinosa shows significant acclimation in the temperature dependence of photosynthesis when grown under different temperature regimes. The presence of leaf hairs which reduce leaf absorptance and consequently leaf temperature plays an important part in the ability of E. farinosa to survive in its native high temperature environment. When the effects of pubescence are taken into account, there are few if any significant differences in the photosynthetic characteristics of the two species.  相似文献   

9.
The endemic Hawaiian species of Scaevola and Euphorbia grow in a wide variety of native habitats and exhibit a wide range of variation in photosynthetic responses. Light-saturated photosynthetic capacities range from 12.0 to 24.7 μmol CO2 m−-2 s−-1 in the Scaevola species and from 18.2 to 51.4 μmol CO2 m−-2 s−-1 in the Euphorbia species. Within each genus, differences in light-saturated photosynthetic capacity are paralleled by differences in mesophyll and leaf conductances to CO2. Within each habitat, the C4 Euphorbia species exhibits a significantly higher photosynthetic capacity and a significantly higher mesophyll conductance than the corresponding C3 Scaevola species. These differences are greatest in the dry scrub habitat and least in the wet forest habitat. One photosynthetic characteristic that exhibits little variation among the species within each genus, yet that exhibits a consistently large difference between the species within each habitat, is photosynthetic water-use efficiency. The C4 Euphorbia species possess water-use efficiencies that are 2–3½ times as high as those of the C3 Scaevola species, regardless of whether these species are native to very dry or very wet habitats. At present, the ecological significance of this large inherent difference in photosynthetic water-use efficiency is unknown. Indeed, it appears that neither photosynthetic pathway has imposed any major inherent constraints on the ability of the Scaevola and Euphorbia species to diversify into a wide variety of habitats.  相似文献   

10.
Summary Common generalizations concerning the ecologic significance of C4 photosynthesis were tested in a study of plant gas exchange, productivity, carbon balance, and water use in monospecific communities of C3 and C4 salt desert shrubs. Contrary to expectations, few of the hypotheses concerning the performance of C4 species were supported. Like the C3 species, Ceratoides lanata, the C4 shrub, Atriplex confertifolia, initiated growth and photosynthetic activity in the cool spring months and also exhibited maximum photosynthetic rates at this time of year. To compete successfully with C3 species, Atriplex may have been forced to evolve the capacity for photosynthesis at low temperatures prevalent during the spring when moisture is most abundant. Maximum photosynthetic rates of Atriplex were lower than those of the C3 species. This was compensated by a prolonged period of low photosynthetic activity in the dry late summer months while Ceratoides became largely inactive. However, the annual photosynthetic carbon fixation per ground area was about the same in these two communities composed of C3 and C4 shrubs. The C4 species did not exhibit greater leaf diffusion resistance than the C3 species. The photosynthesis/transpiration ratios of the two species were about the same during the period of maximum photosynthetic rates in the spring. During the warm summer months the C4 species did have superior photosynthesis/transpiration ratios. Yet, since Ceratoides completed a somewhat greater proportion of its annual carbon fixation earlier in the season, the ratio of annual carbon fixation/transpiratory water loss in the two communities was about the same. Atriplex did incorporate a greater percentage of the annual carbon fixation into biomass production than did Ceratoides. However, this is considered to be a reflection of properties apart from the C4 photosynthetic pathway. Both species displayed a heavy commitment of carbon to the belowground system, and only about half of the annual moisture resource was utilized in both communities.  相似文献   

11.
The non-graminaceous wild flora of Hungary was screened for C4 plants by using the stable carbon isotope ratio, the leaf anatomy and the photosynthetic carbon dioxide compensation concentration to determine the photosynthetic pathway type. On the whole, 31 C4 species (native or naturalized) were found in the Amaranthaceae, Chenopodiaceae, Cyperaceae, Euphorbiaceae, Portulacaceae and Zygophyllaceae families. Together with the 26 C4 grass species (Poaceae) reported earlier (Kalapos 1991), a total of 57 wild C4 species occur in Hungary, which forms 2.6 % of the country's angiosperm flora. This figure is somewhat higher than what was expected on climatic grounds, a fact probably due to certain edaphic conditions favouring C4 plant growth. In Hungary, the C4 species are predominantly annuals growing in open habitats such as dry grasslands, inland saline areas, temporarily exposed riverbeds and disturbed sites. In comparison with C3 plants, the C4 species have higher temperature and light preferences, and their phenology lags behind that of the C3 plants. These differences might account for C4 plants being usually excluded from productive biotopes in Hungary, where the C3 canopy may become closed during the growing season before C4 plants can start their ontogenetic development. Ecological properties of C3 and C4 plants differ considerably in the Cyperaceae, but much less in the Chenopodianceae family. Among C4 annuals naturalized aliens are common, most of which colonized hungary in the last two centuries. Increasing preponderance of C4 plants is anticipated in the future as a consequence of possible climate changes and the ever increasing human impact on terrestrial vegetation. This revised version was published online in August 2006 with corrections to the Cover Date.  相似文献   

12.
Summary Diurnal patterns of CO2 exchange and titratable acidity were monitored in six species of evergreen rosette plants growing in controlled environment chambers and under outdoor environmental conditions. These patterns indicated that two of the species, Yucca baccata and Y. torreyi, were constituitive CAM plants while the other species, Y. elata, Y. campestris, Nolina microcarpa and Dasylirion wheeleri, were C3 plants. The C3 species did not exhibit CAM when grown in any of several different temperature, photoperiod, and moisture regimes. Both photosynthetic pathway types appear adapted to desert environments and all species show environmentally induced changes in their photosynthetic responses consistent with desert adaptation. The results of this study do not indicate that changes in the photosynthetic pathway type are an adaptation in any of these species.  相似文献   

13.
Ecotypic differences in the photosynthetic carbon metabolism of Mollugo verticillata were studied. Variations in C3 and C4 cycle activity are apparently due to differences in the activities of enzymes associated with each pathway. Compared to C4 plants, the activities of C4 pathway enzymes were generally lower in M. verticillata, with the exception of the decarboxylase enzyme, NAD malic enzyme. The combined total carboxylase enzyme activity of M. verticillata was greater than that of C3 plants, possibly accounting for the high photosynthetic rates of this species. Unlike either C3 or C4 plants, ribulose bisphosphate carboxylase was present in both mesophyll and bundle sheath cell chloroplasts in M. verticillata. The localization of this enzyme in both cells in this plant, in conjunction with an efficient C4 acid decarboxylation mechanism most likely localized in bundle sheath cell mitochondria, may account for intermediate photorespiration levels previously observed in this species.  相似文献   

14.
Wang  R.Z. 《Photosynthetica》2002,40(1):97-102
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.  相似文献   

15.
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.  相似文献   

16.
Summary Mono-specific communities of the C3 sedge, Scirpus olneyi and the C4 grass, Spartina patens, were exposed to normal ambient or elevated CO2, (ca. 680 l l–1) throughout the 1987 and 1988 growing seasons in open-top field chambers located on a tidal marsh. Single stems of C3 plants grown in ambient or elevated CO2 showed an increased photosynthetic rate when tested at elevated CO2 for both seasons. This increase in photosynthetic response in the C3 species was maintained throughout the 1987 and 1988 growing season. The stimulation of photosynthesis with elevated CO2 appeared to increase as temperature increased and decreased as photosynthetic photon flux (PPF) increased. Analysis of the photosynthetic response of the C3 species during the 1988 season indicated that significant differences in light-saturated photosynthetic rate between ambient and elevated CO2 conditions continued until October. In contrast to the C3 sedge, the C4 grass showed no significant photosynthetic increase to elevated CO2 except at the beginning of the 1988 season.  相似文献   

17.
Question: Are trait differences between grasses along a gradient related to climatic variables and/or photosynthetic pathway? Location: Temperate grassland areas of South and North America. Methods: In a common garden experiment, we cultivated C3 and C4 grasses from grasslands under different climatic conditions, and we measured a set of 12 plant traits related to size and resource capture and utilization. We described (1) interspecific plant trait differences along a climatic gradient defined by the precipitation and temperature at the location where each species is dominant and (2) the association between those plant trait differences and the photosynthetic pathway of the species. Results: Trait differences between grasses were related to the precipitation at the area where each species is dominant, and to the photosynthetic pathway of the species. Leaf length, leaf width, plant height, leaf area per tiller, specific leaf area, leaf δ13C ratio, and nitrogen resorption efficiency increased while leaf dry matter content and nitrogen concentration in senesced leaves decreased as precipitation increased. A proportion of these changes along the gradient was related to the photosynthetic pathway because dominant grass species in cold areas with low precipitation are mainly C3 and those from warm and wet areas are C4. Conclusions: A previous worldwide analysis showed that traits of graminoid species measured in situ changed slightly along climatic gradients (< 10% variance explained). In contrast, under a common environment we observed that (1) grass traits changed strongly along a climatic gradient (30‐85% variance explained) and, (2) a proportion of those changes were related to the association between photosynthetic pathway of the species and precipitation.  相似文献   

18.
C3 desert plant Reaumuria soongorica (RS-C3) and C4 desert plant Salsola passerina (SP-C4) may exist either in individual or in associated communities. Carbon isotope composition, leaf water potential, gas exchange and chlorophyll fluorescence characteristics of the individual and associated communities were compared with reveal, whether the associated growth represent an advantage under harsh habitat. The results showed that the ??13C values of leaves of RS-C3 and SP-C4 across different habitats fluctuated, respectively, from ?24 to ?27??? and from ?14 to ?16???. Leaf water potential of RS-C3 was lower than SP-C4 all day long, growing either individually or associated with the C3 plant. When associated with the C4 plant, the net photosynthetic rate of the RS-C3 increased, and the photosynthetic rate of the partner SP-C4 decreased. The transpiration rates of the associated RS-C3 and SP-C4 were both lower than in their individual colonies. In associated communities, in RS-C3, the maximal photochemical efficiency, the effective photochemical efficiency, the relative electron transport rate, the photochemical quenching of PS II increased, and the non-photochemical quenching of PS II decreased; all these parameters changed oppositely in the SP-C4 plant. This shows that, in the associated community, the C4 plants might facilitate adaptation of the RS-C3, while SP-C4 plant can adapt to the harsh environment through their own specialties. The association favored the expression of natural photosynthetic characteristics and survival of RS-C3, while retarded the growth of SP-C4. Associated growth decreases the transpiration rate of the whole community; it is conducive to improve its water use efficiency.  相似文献   

19.
Wang  R.Z. 《Photosynthetica》2004,42(2):219-227
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.  相似文献   

20.
Cyanobacteria are environmentally important photosynthetic microorganisms attracting a growing attention in various areas of basic and applied researches. To better understand their metabolism, we presently report on the development of a robust and simple protocol for facile extraction and high throughput analysis of the metabolites of the widely-used strain Synechocystis PCC6803 through liquid chromatography coupled to high resolution mass spectrometry (LC/MS). Our analytical method was developed and tested with 102 reference compounds representative of the chemical diversity of polar cell metabolites, and Synechocystis cell extracts spiked with 37 reference compounds. These samples were analyzed with two chromatographic systems, each coupled to a LTQ-Orbitrap mass spectrometer: a liquid chromatographic system equipped with a pentafluorophenylpropyl column (the PFPP-LC/MS system), and an ultra-high performance liquid chromatographic system with a C18-reversed phase column (the C18-UHPLC/MS system). We showed that the PFPP-LC/MS method performs better than the C18-UHPLC/MS method in terms of retention, separation and detection of metabolites. Consequently, we applied the PFPP-LC/MS method to analyze the metabolome of Synechocystis growing under various conditions of light and glucose, which strongly influence cell growth. We found that glucose increases glucose storage (synthesis of glycogen-like polysaccharide) and catabolism (oxidative pentose phosphate pathway and glycolysis), while it decreases the Calvin–Benson cycle that consumes photosynthetic electrons for CO2 assimilation. Depending on light and glucose availabilities, this global metabolic reprogramming can generate an oxidative stress, likely through the recombination of the glucose-spared electrons with the photosynthetic oxygen thereby producing toxic reactive oxygen species.  相似文献   

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