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.     

Carbon isotope ratio measurements of succulent plants in southern Africa

Summary Succulent plants representing 16 families and a variety of growth forms originating from winter, summer, and year-round rainfall regimes in southern Africa were analyzed for carbon isotope ratios. Most families had species with ¹³C values indicative of CAM, particularly those from winter and year-round rainfall regimes. Plants with ¹³C values intermediate between CAM and C₃, indicating flexible photosynthetic pathways, were generally leafy perennials subject to seasonal tissue dehydration. Reproductive tissue tended to have less negative ¹³C values than vegetative tissue on the same plant, indicating drought-season origin of the former.CIW-DPB Publication No. 609     

Characterization of photosynthetic pathway of plant species growing in the eastern Tibetan plateau using stable carbon isotope composition

The photosynthetic pathway of plant species collected at Menyuan, Henan, and Maduo sites, east of Tibetan Plateau, China, during the growing season were studied using stable carbon isotopes in leaves. The 232 samples leaves analyzed belonged to 161 species, 30 families, and 94 genera. The δ¹³C values (from −24.6 to − 29.2 ‰) indicated that all the considered species had a photosynthetic C₃ pathway. The absence of plant species with C₄ photosynthetic pathway might be due to the extremely low air temperature characterizing the Tibetan Plateau. The average δ¹³C value was significantly (p<0.05) different between annuals and perennials at the three considered study sites. Hence the longer-lived species had greater water-use efficiency (WUE) than shorter-lived species, that is, longer-lived species are better adapted to the extreme environmental conditions of the Tibetan Plateau.     

Comparative ecophysiology of five species of Sedum (Crassulaceae) under well-watered and drought-stressed conditions

Summary Gas exchange patterns, diurnal malic acid fluctuations, and stable carbon isotope ratios of five species of Sedum were investigated to assess the ecophysiological characteristics of three different photosynthetic pathways under well-watered and drought-stressed conditions. All five species have succulent leaves and stems and were examined under identical environmental conditions. When well-watered, Sedum integrifolium (Raf.) Nels. and S. ternatum Michx. displayed C₃ photosynthesis, S. telephioides Michx. and S. nuttallianum Raf. exhibited CAM-cycling, and S. wrightii A. Gray showed CAM. When grown under a less frequent watering regime, S. integrifolium and S. ternatum exhibited CAM-cycling, whereas S. telephioides and S. nuttallianum displayed CAM-cycling simultaneously with low-level CAM. Sedum wrightii retained its CAM mode of photosynthesis. In general, leaf ¹³C values reflected these variations in photosynthetic pathways. While all values of water-use efficiency (WUE) were greater than those reported for most C₃ and C₄ species, no correlation of malic acid accumulation in the CAM and CAM-cycling (including low-level CAM) species with increased WUE was found. Sedum wrightii (CAM) had the highest WUE value at night, yet its 24-h WUE was not different from S. ternatum when the latter was in the C₃ mode. Thus, relative water-use efficiencies of these species of Sedum were not predictable based on photosynthetic pathways alone.     

The effects of salinity,crassulacean acid metabolism and plant age on the carbon isotope composition of <Emphasis Type="Italic">Mesembryanthemum crystallinum</Emphasis> L., a halophytic C<Subscript>3</Subscript>-CAM species

The carbon isotope composition of the halophyte Mesembryanthemum crystallinum L. (Aizoaceae) changes when plants are exposed to environmental stress and when they shift from C₃ to crassulacean acid metabolism (CAM). We examined the coupling between carbon isotope composition and photosynthetic pathway by subjecting plants of different ages to salinity and humidity treatments. Whole shoot ¹³C values became less negative in plants that were exposed to 400 mM NaCl in the hydroponic solution. The isotopic change had two components: a direct NaCl effect that was greatest in plants still operating in the C₃ mode and decreased proportionally with increasing levels of dark fixation, and a second component related to the degree of CAM expression. Ignoring the presumably diffusion-related NaCl effect on carbon isotope ratios results in an overestimation of nocturnal CO₂ gain in comparison to an isotope versus nocturnal CO₂ gain calibration established previously for C₃ and CAM species grown under well-watered conditions. It is widely taken for granted that the shift to CAM in M. crystallinum is partially under developmental control and that CAM is inevitably expressed in mature plants. Plants, cultivated under non-saline conditions and high relative humidity (RH) for up to 63 days, maintained diel CO₂ gas-exchange patterns and ¹³C values typical of C₃ plants. However, a weak CAM gas-exchange pattern and an increase in ¹³C value were observed in non-salt-treated plants grown at reduced RH. These observations are consistent with environmental control rather than developmental control of the induction of CAM in mature M. crystallinum under non-saline conditions.     

Stable carbon isotope characteristics of desert plants in the Junggar Basin, China

Desert plants have unique strategies for survival and growth to cope with the limited water availability in arid regions. The stable carbon isotope (δ ¹³C) provides an integrated measurement of internal plant physiological and external environmental properties affecting photosynthetic gas exchange and water use efficiency. The δ ¹³C values of 84 species in the Junggar Basin were categorized into two groups (ranged from −30.1 to −23.3‰ for C₃ and −14.9 to −9.9‰ for C₄ species, respectively). No life form differences in δ ¹³C values were detected in C₃ (p = 0.78) and C₄ plants (p = 0.63). Small differences among life forms were observed in δ ¹³C values in C₄ species with shrubs slightly depleted (−13.3‰) relative to perennials (−13.1‰) and annuals (−12.5‰). These differences suggested that δ ¹³C value could not represent a plant functional group classification based on life forms in C₄ plants in extremely arid regions. Ephemerals are all using C₃ photosynthetic pathway and no significant differences (p = 0.92) in δ ¹³C values were observed between annuals (−26.5‰) and perennials (−26.4‰). The δ ¹³C values of Tulipa iliensis (an important ephemeral species distributed widely in the Junggar Basin) among nine natural populations were positively correlated with leaf (r ² = 0.46, p = 0.046) and soil (r ² = 0.67, p = 0.007) total nitrogen content, and negatively correlated with leaf (r ² = 0.48, p = 0.039) and soil (r ² = 0.79, p = 0.001) water content. This indicated that the variation in δ ¹³C values of T. iliensis was probably caused by both water availability associated stomatal openness and nitrogen availability associated photosynthetic capacity. T. iliensis is very sensitive to water and nitrogen availability in soil.     

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.     

Seasonal response to drought and rewatering in Portulacaria afra (L.) Jacq.

Summary Gas exchange characteristics of droughted and rewatered Portulacaria afra were studied during the seasonal shift from CAM to C₃ photosynthesis. ¹⁴CO₂ uptake, stomatal conductance, and total titratable acidity were determined for both irrigated and 2, 4, and 7.5 month waterstressed plants from summer 1984 to summer 1985. Irrigated P. afra plants were utilizing the CAM pathway throughout the summer and shifted to C₃ during the winter and spring. Beginning in September, P. afra plants shifted from CAM to CAM-idling after 2 months of water-stress. When water-stress was initiated later in the fall, exogenous CO₂ uptake was still measurable after 4 months of drought. After 7.5 months of stress, exogenous CO₂ uptake was absent. The shift from CAM to CAM-idling or C₃ in the fall and winter was related to when water stress was initiated and not to the duration of the stress. Gas exchange resumed within 24 h of rewatering regardless of the duration of the drought. In the winter and spring, rewatering resulted in a full resumption of daytime CO₂ uptake. Whereas during the summer, rewatering quickly resulted in early morning CO₂ uptake, but nocturnal CO₂ uptake through the CAM pathway was observed after 7 days. Gas exchange measurements, rewatering characteristics, and transpirational water loss support the hypothesis that the C₃ pathway was favored during the winter and spring. The CAM pathway was functional during the summer when potential for water loss was greater. Our investigations indicate that P. afra has a flexible photosynthetic system that can withstand long-term drought and has a rapid response to rewatering.     

Ecological evidence concerning the adaptive significance of the C4 dicarboxylic acid pathway of photosynthesis

Summary Ecological data were analyzed to determine the environmental conditions associated with the occurrence of plants possessing C₃ and C₄ photosynthetic pathways. Non-parametric analysis of variance, multiple regression analysis and discriminant analysis techniques were applied to information derived from separate studies on plant species in California and central Europe. All the analyses revealed significant differences in ecology between the C₃ and C₄ groups. The occurrence of C₄ species was greater where summer or winter temperatures were relatively high and moisture availability relatively low. Using discriminant analysis, more than 80% of the species considered were classified into the correct photosynthetic category on the basis of their distribution with respect to temperature. No significant differences between the C₃ and C₄ groups were found for other environmental factors, including light, soil nitrogen, soil salinity and continentality of climate. These results support the suggestion from previous physiological studies that C₄ species may possess competitive advantages under conditions of high temperature and intermittent water stress. The C₄ species were relatively restricted in the range of environmental conditions where they occurred, and as a group C₄ species may be ecologically more specialized than C₃ species.     

Geographical and environmental distribution of C3 and C4 grasses in the Sinai,Negev, and Judean deserts

Summary The relation between photosynthetic pathway and habitat of the grass species recorded in the desert regions of Sinai, Negev, and Judea was investigated. The climatic conditions and micro-environments in the study area vary considerably, and the distribution of the various species is found to conform to specific patterns which reveal the adaptive advantages of the different photosynthetic pathways. There is also a distinct correlation between the phytogeographic origin of the grass species and the photosynthetic pathways that they utilize.The survey shows that the majority of the grass species in the region are of the C₃ type and all except one of these species belong to the Holarctic domain. This is in accordance with the fact that the region forms part of the Mediterranean winter rainfall regime and that C₃ species have an adaptive advantage where minimum temperatures are low during the winter growing season.The occurence of C₄ species increases with decreasing rainfall and they dominate in those districts where temperatures are high throughout the year. These C₄ grasses are of both Holarctic and Palaeotropic origin according to the classification adopted here, but they are essentially all elements of the Saharo-Arabian, Irano-Turanian, Sudanian, or Tropical phytogeographic regions and are not typical of the Mediterranean or Euro-Siberian floras. The plants with multi-regional distributions that occur in Mediterranean communities may well be intrusive.Analysis of the three subtypes of the C₄ species suggests that the malate-forming NADP-me grasses grow where water stress is not a dominating factor, while the aspartateforming NAD-me grasses are more successful under xeric conditions. The PEP-ck species are not abundant and form an intermediate group between the NADP-me and NAD-me subtypes.     

The photosynthetic pathway types of some desert plants from India,Saudi Arabia,Egypt, and Iraq

Summary Plants collected from different habitats in the deserts of India, Saudi Arabia, Egypt, and Iraq were screened for their photosynthetic CO₂-fixation pathways using ¹³C and D values. The analyses comprised 128 species belonging to 108 genera and 46 families. Neither the C₄ nor the CAM pathway was prevalent in the plant families analyzed except in Poaceae, where C₄ metabolism absolutely dominated. Of 93 dicotyledonous plants, only 10 exhibited a C₄ pathway and only 2 were CAM plants. The study shows that some species reported by other workers as C₄ plants are clearly C₃ ones.Dedicated to Prof. Dr. Hermann Merxmüller on the occasion of his 60th birthday     

Restriction analysis and sequencing of the ITS regions and 5.8S gene of rDNA of Ganoderma isolates from infected oil palm and coconut stumps in Malaysia

Use of Crassulacean acid metabolism (CAM) plants in México and worldwide has a long history, but the morphological and photosynthetic aspects of these plants have only been considered recently. Emphasis in this article is on the daily net CO₂ uptake ability by three species of agaves and three species of cacti that are currently extensively cultivated in México for beverages, food, fodder, and forage ‐ Agave mapisaga, A. salmiana, A. tequilana, Opuntia ficus‐indica, O. robusta and Stenocereus queretaroensis. Data under controlled conditions are used to help interpret seasonal net CO₂ uptake patterns observed in the field. These CAM plants have instantaneous and total daily net CO₂ uptake values similar to those for highly productive C₃ and C₄ crops. The future increase in the cultivated area of CAM plants will have both agronomical and ecological ramifications because of the ability of these plants to endure prolonged drought and to sequester carbon during extended dry periods when few C₃ and C₄ crops and non‐CAM native plants can fix atmospheric CO₂.     

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.     

Photosynthetic pathways inChenopodiaceae from Africa, Asia and Europe with their ecological, phytogeographical and taxonomical importance

Photosynthetic pathways are reported for 305 species ofChenopodiaceae from Africa, Asia and Europe. Ecological characteristics, phytogeography and life forms of all species are given, and their correlation with relevant CO₂-fixation types are discussed. 205 species (67.2%) exhibit the C₄-pathway and 100 species (32.8%) the C₃-pathway of CO₂-fixation. Most of the C₄ species are of Irano-Turanian origin. The diversity of C₄ species of the Irano-Turanian phytochoria, with very harsh winters, are interpreted by the active period of Chenopods in summer. There is a close relationship between some special morphotypes and respective photosynthetic type. Halophytes and xerophytes with articulated stems and stem succulents ofAnabasis-type are exclusively C₄. Leaf succulent halophytes and xerohalophytes are predominantly C₄. Hygrohalophytes with leaf or stem succulence are often C₃. Probably many C₄ Salsoleae have been evolved in the Afroasiatic arid zone after the climatic changes of the Miocene. Among them there is a high proportion of annuals that have a younger origin. There is some evidence that the present Chenopods of the subfamily Salsoloideae may have their ancient stock in NW Africa. We found close taxonomic relationships between photosynthetic pathways and infrageneric classification in genera with both types of C₃ and C₄, likeAtriplex andSuaeda. Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer, Wien, on the occasion of his 70th birthday     

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.     

Occurrence and ecological characteristics of C4 dicot and Cyperaceae species in the Hungarian flora

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.     

The small scale spatial pattern of C3 and C4 grasses depends on shrub distribution

At micro‐site scale, the spatial pattern of a plant species depends on several factors including interactions with neighbours. It has been seen that unfavourable effects generate a negative association between plants, while beneficial effects generate a positive association. In grasslands, the presence of shrubby species promotes a particular microenvironment beneath their canopy that could affect differently the spatial distribution of plants with different tolerance to abiotic conditions. We measured photosynthetic active radiation, air temperature and wind speed under shrub canopies and in adjacent open sites and analysed the spatial distribution of four grass species (two C₃ and two C₄) in relation to shrub canopy in a grazed sub‐humid natural grassland in southern Uruguay. Radiation, air temperature and wind speed were lower under shrubs than in adjacent open sites. The spatial distribution of grasses relative to the shrub canopy varied depending on the photosynthetic metabolism of grasses. C₄ grasses showed a negative association or no correlation with the shrubs, whereas C₃ grasses showed a positive association. Our results highlight the importance of the photosynthetic metabolism of the grasses in the final outcome of interactions between grasses and shrubs. Micro‐environmental conditions generated underneath shrubs create a more suitable site for the establishment of C₃ than for C₄ grasses. These results show that facilitation could be more important than previously thought in sub‐humid grasslands.     

Phosotynthesis in hemiepiphytic species of Clusia and Ficus

Summary Hemiepiphytic species in the genera Clusia and Ficus were investigated to study their mode of photosynthetic metabolism when growing under natural conditions. Despite growing sympatrically in many areas and having the same growth habit, some Clusia species show Crassulacean acid metabolism (CAM) whereas all species of Ficus investigated are C₃. This conclusion is based on diurnal CO₂ fixation patterns, diurnal stomatal conductances, diurnal titratable acidity fluctuations, and ¹³C isotope ratios. Clusia minor, growing in the savannas adjacent to Barinas, Venezuela, shows all aspects of Crassulacean acid metabolism (CAM) on the basis of nocturnal gas exchange, stomatal conductance, total titratable acidity, and carbon isotope composition when measured during the dry season (February 1986). During the wet season (June 1986), the plants shifted to C₃-type gas exchange with all CO₂ uptake occurring during the daylight hours. The carbon isotope composition of new growth was-28 to-29 typical of C₃ plants.     

GERMINATION ECOPHYSIOLOGY OF HERBACEOUS PLANT SPECIES IN A TEMPERATE REGION

Germination phenology data have been collected from 75 winter annuals, 49 summer annuals, 28 monocarpic perennials, and 122 polycarpic perennials, and experimental investigations of dormancy breaking and germination requirements have been conducted on 56 winter annuals, 32 summer annuals, 18 monocarpic perennials, and 73 polycarpic perennials. The purpose of these studies was to determine if there are correlations between the dormancy breaking and germination requirements of seeds and the germination phenology, life cycle type, habitat requirements, range of geographical distribution, and phylogenetic relationships of the species. Germination phenology is highly correlated with the responses of seeds to the yearly temperature cycle. Species with winter and summer annual life cycles have predictable germination characteristics, but monocarpic and polycarpic perennials do not. Several dormancy types may be found in a given habitat, and narrowly endemic and widely-distributed species in the same genus may have similar germination characteristics. Within some families there is a tendency for a particular type of seed-temperature response to be very important, but frequently this is related to the predominance of a given life cycle type in the family.     

C4 photosynthesis in the vegetation of Aldabra Atoll

Summary The occurrence of the C₄ photosynthetic system, based upon an examination of leaf anatomy, of nearly all species of the vegetation of Aldabra Atoll, western Indian Ocean, is reported. About 19% of the flora has the C₄ system. It only occurs in herbaceous plants, which grow either in grasslands or as an understorey to tall shrubs. Both C₃ and C₄ species compete side by side in the mixed-species communities.The Royal Society Aldabra Research Station