Vegetation variation within and among palustrine wetlands along an altitudinal gradient in KwaZulu-Natal,South Africa

The variation in graminoid species composition and diversity and the distribution of photosynthetic pathways among 66 wetlands in KwaZulu-Natal, South Africa, and within six of these wetlands was described and related to measured physical parameters, using multivariate and univariate techniques. Altitude, which ranged from 550 m to 2120 m, accounted for most variation among wetlands, with an almost complete turnover of species along this gradient. Landform setting was less important in explaining overall species composition, but relationships of individual species were revealed (e.g. Eleocharis dregeana showed an affinity for depressions). Within a wetland there was an almost complete turnover of species along a gradient of wetness, as described using soil morphological criteria. Most species were consistently associated with the same wetness zones across different wetland sites, e.g., Phragmites australis with the wettest zone, Pycreus macranthus with the intermediate zone, and Eragrostis plana with the least wet zone. The occurrence and abundance of different photosynthetic pathway types depended on altitude and degree of wetness. At high altitudes, C₃ sedges, notably Carex acutiformis, dominated the wettest zone and C₃ and C₄ grasses and sedges dominated the intermediate and least wet zones. At mid altitudes, C₃ and C₄ sedges and C₃ grasses dominated the wettest zone, C₃ and C₄ grasses and sedges dominated the intermediate zone and C₄ grasses dominated the least wet zone. Low altitude sites showed a similar distribution of photosynthetic pathways as mid-altitude sites, but C₃ species were less abundant. Species richness was positively associated with the log of wetland size and, at the level of an individual wetland, species richness and evenness were found to be consistently greater in the intermediate and least wet zone compared with the wettest zone. The management implications of the results are discussed in the light of continuing anthropogenic loss of wetlands in the study area and global climate change.     

Changes in individual plant traits and biomass allocation in alpine meadow with elevation variation on the Qinghai-Tibetan Plateau

Plant traits and individual plant biomass allocation of 57 perennial herbaceous species, belonging to three common functional groups (forbs, grasses and sedges) at subalpine (3700 m ASL), alpine (4300 m ASL) and subnival (⩾5000 m ASL) sites were examined to test the hypothesis that at high altitudes, plants reduce the proportion of aboveground parts and allocate more biomass to belowground parts, especially storage organs, as altitude increases, so as to geminate and resist environmental stress. However, results indicate that some divergence in biomass allocation exists among organs. With increasing altitude, the mean fractions of total biomass allocated to aboveground parts decreased. The mean fractions of total biomass allocation to storage organs at the subalpine site (7%±2% S.E.) were distinct from those at the alpine (23%±6%) and subnival (21%±6%) sites, while the proportions of green leaves at all altitudes remained almost constant. At 4300 m and 5000 m, the mean fractions of flower stems decreased by 45% and 41%, respectively, while fine roots increased by 86% and 102%, respectively. Specific leaf areas and leaf areas of forbs and grasses deceased with rising elevation, while sedges showed opposite trends. For all three functional groups, leaf area ratio and leaf area root mass ratio decreased, while fine root biomass increased at higher altitudes. Biomass allocation patterns of alpine plants were characterized by a reduction in aboveground reproductive organs and enlargement of fine roots, while the proportion of leaves remained stable. It was beneficial for high altitude plants to compensate carbon gain and nutrient uptake under low temperature and limited nutrients by stabilizing biomass investment to photosynthetic structures and increasing the absorption surface area of fine roots. In contrast to forbs and grasses that had high mycorrhizal infection, sedges had higher single leaf area and more root fraction, especially fine roots.     

The distribution of C3 and C4 grasses and carbon isotope discrimination along an altitudinal and moisture gradient in Kenya

Summary More than 500 species of the Poaceae are found in Kenya, East Africa. Eighteen of twenty-seven tribes are exclusively (except the Paniceae and Danthonieae) of the C₃ photosynthetic type. A floristic analysis of low altitude grasslands suggests that nearly all species at these low altitudes are of the C₄ photosynthetic type. At high altitudes, however, nearly all grasses are of the C₃ photosynthetic type. Open grassland vegetation was sampled along a transect from arid low altitude sites to the top of Mt. Kenya in an attempt to document the general distributions of the photosynthetic types.The major tribes illustrated three general patterns of distribution. The C₄ tribes Chlorideae, Eragrosteae, Sporoboleae, and Aristideae were abundant at low altitudes (or low indices of available soil moisture). The Paniceae and Andropogoneae were also exclusively C₄ but were more common at intermediate altitudes. The C₃ tribes Aveneae, Festuceae, and Agrostideae were found only at high altitudes. In these open grasslands there were no C₃ species below 2,000 m and no C₄ species above 3,000 m. The variation in ₁₃C of the live grass vegetation with altitude confirms these distributional patterns and suggests a sharp transition zone between these two photosynthetic types. The photosynthetic type accounts for broad distributions within the Poaceae but these distributions are further modified by characteristics which may be inherent in the tribal groups. Ecological and paleoecological significance of these patterns of distribution are discussed.     

Geographical distribution and chorology of grasses in the Arabian Peninsula

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

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.     

Silica Deposition in Some C3 and C4 Species of Grasses, Sedges and Composites in the USA

We report new information on silica deposition in 15 plant species,including nine grasses, two sedges and four composites. Thesilica depositional patterns found in seven of the grass speciesindicate that they are C₄ plants. However the festucoid grassCortaderia selloana is a C₃ plant with long leaf trichomes andoval silica structures in the leaves. In contrast the panicoidC₄ grasses Chasmathium latifolium, Chasmathium sessiflorum,Imperata cylindrica, Panicum repens, Panicum commutatum andSetaria magna, all produce dumb-bell-shaped silica structuresin the leaves. The chloridoid grasses Spartina patens and Spartinacynosuroides have saddle-shaped structures and no dumb-bellor oval shaped ones. The sedges Rhynchospora plumosa and Scirpuscyperinus were found to have oval phytoliths and may be C₃ plants.Our examination of these and other grasses strongly suggeststhat C₄ grasses tend to produce the same type of silica cells.Grasses and sedges with C₃ type photosynthesis tend to produceoval silica structures. The composite Grindelia squarrosa andsunflowers Helianthus angustifolia, Helianthus atrorubens andHelianthus tuberosus absorb relatively small amounts of siliconand larger amounts of calcium, where both elements deposit inleaf trichomes. We found no clear indicator for the C₃ sunflowersor C₄ types in the Asteraceae. Helianthus tuberosus leaves havemany trichomes on the adaxial surface. These trichomes havea higher concentration of silica than the surrounding leaf surface.Helianthus tuberosus leaves had much higher ash and silica contentsthan those of Helianthus angustifolia and Helianthus atrorubens.The composite Grindelia squarrosa has a usual deposition ofsilica in the basal cells around the guard cells. Silica depositionoften reflects the surface features of a leaf. An exceptionis Scripus cyperinus where the silica structures are deep inthe tissue and do not reflect the surface configurations. Theinforescence of Setaria magna had a 14.64 silica content. Thetufts of white, silky hairs characteristic of Imperata cylindricainflorescence have no silica. C₃ and C₄ plants, silica and ash content, scanning electron microscopy, energy-dispersive X-ray analysis, silicon distribution, spectra of elements in plants, trichomes, silica fibres, phytoliths     

Aboveground Biomass Removal by Burning and Raking Increases Diversity in a Reconstructed Prairie

Prescribed spring burning often contributes to a predominance of C₄ grasses and low forb abundance and is impractical at many sites, especially near development. We tested raking after mowing as an alternative to prescribed burning in a reconstructed Minnesota prairie. We also tested mowing without raking as a possible means of maintaining prairie communities. Frequency, flowering stem abundance, and cover were measured for all plant species and native functional groups (C₄ grasses, C₃ graminoids, forbs, legumes, and annual or biennial forbs). Mowing alone did not differ from the control in its effect on any functional groups of plants. Round‐headed bush clover (Lespedeza capitata), a legume, and Black‐eyed Susan (Rudbeckia hirta), a biennial, increased in frequency with treatments that removed biomass (i.e., fire or raking), but they did not have significantly more flowering stems. Thus, new plants established well from seed, whereas the vitality of mature plants did not change. Raking had similar effects to burning on most functional groups, although flowering stems of C₄ grasses were significantly more abundant after fire than after raking. Burning reduced some C₃ forbs and grasses and favored the dominance of C₄ grasses. Therefore, raking after mowing in the spring provides an alternative to prescribed burning that has many of the same positive aspects as fire but does not promote aggressive C₄ grasses to the same extent.     

Effects of charring on the carbon isotopic composition of grass (Poaceae) epidermis

Charred modern grass epidermis preserves the carbon isotopic composition of the parent plant photosynthetic pathway. Fifty-nine modern grasses and sedges were collected in lowland western Uganda. All charred epidermal samples from C₄ grasses or sedges preserve a carbon isotopic value within the range typical for C₄ plants (−17 to −10‰), and charred epidermal fragments from C₃ plants have carbon isotopic values between −30 and −26‰. The process of charring results in a slightly enriched carbon isotopic signature (−11.9‰ mean charred value as compared to −12.8‰ mean unaltered grass tissue value). δ¹³C measurements of replicate samples from the same plant vary within 1–2‰, yet all values for the same plant stay within the expected values for the photosynthetic pathway of the plant. δ¹³C measurements on >180-μm charred grass epidermal fragments extracted from surface sediment samples from three lakes on the lowland western Ugandan landscape confirm the predominant lowland C₄ grass input (δ¹³C=−16 to −19‰). These results demonstrate the utility of using carbon isotopic analysis of charred grass epidermis to reconstruct C₃ vs. C₄ grassland assemblages on the landscape. Furthermore, such downcore δ¹³C profiles can be used to highlight key zones of C₃ vs. C₄ grass change for which taxonomic analysis of fossil grass epidermis could provide more detailed information regarding grassland community composition.     

C4 photosynthesis evolved in warm climates but promoted migration to cooler ones

C₄ photosynthesis is considered an adaptation to warm climates, where its functional benefits are greatest and C₄ plants achieve their highest diversity and dominance. However, whether inherent physiological barriers impede the persistence of C₄ species in cool environments remains debated. Here, we use large grass phylogenetic and geographical distribution data sets to test whether (1) temperature influences the rate of C₄ origins, (2) photosynthetic types affect the rate of migration among climatic zones, and (3) C₄ evolution changes the breadth of the temperature niche. Our analyses show that C₄ photosynthesis in grasses originated in tropical climates, and that C₃ grasses were more likely to colonise cold climates. However, migration rates among tropical and temperate climates were higher in C₄ grasses. Therefore, while the origins of C₄ photosynthesis were concentrated in tropical climates, its physiological benefits across a broad temperature range expanded the niche into warmer climates and enabled diversification into cooler environments.     

Grassland development under glacial and interglacial conditions in southern Africa: review of pollen,phytolith and isotope evidence

Pollen evidence suggests that grasslands were well established in southern Africa by the Late Tertiary. Evidence for grassland composition in the region during the Quaternary includes published accounts of isotopes, grass phytoliths and pollen of both grasses and woody plants from a wide range of different environments. Isotope data were derived from speleothems (stalagmites), fossil bones, and fossil tooth enamel and plant material in fossil hyrax dung. The different data types suggest that, with perhaps the exception of the dry southern Kalahari region, temperate grassland consisted of a relatively increased C₃- to C₄-grass ratios during the Last Glacial Maximum (LGM). Cold winter temperature extremes in the southern high latitude and altitude regions and a persistent winter rainfall pattern over the Cape region during the LGM probably limited the distribution of C₄ grasses and canceled out any advantages gained from lowered CO₂ concentrations in the atmosphere. In contrast, in the tropics where marked seasonal temperature fluctuations were lacking, C₄-grass growth was favored.     

The food web of a severely contaminated site following reclamation with warm season grasses

We used the stable isotope ¹³C to distinguish between food web components that depended on warm season grasses with the C₄ photosynthetic pathway and those that depended on plants with the C₃ pathway. The study site was contaminated by heavy metals from a zinc smelter that operated near Palmerton, Pennsylvania, U.S.A. C₃ plants only contributed 1.16% of aboveground primary productivity, whereas recently seeded (5–7 year old) warm season C₄ grasses contributed the remaining 98.84%. Analyses of tissue samples revealed that the carbon content of invertebrates and vertebrates did not reflect the composition of the vegetation. Of 135 samples, 48 (36%) had greater than 75% of their carbon from C₄‐derived sources, while 32 (24%) of the samples had less than 25%. However, carbon from C₄ grasses passed through to higher trophic levels, as shown by the abundance of predators with a high proportion of C₄‐derived carbon. We document three channels of carbon flux through the food web, one based on warm season grasses, now supporting a functioning ecosystem at all key trophic levels, one based on C₃ plants, and a third based on detritus. Theoretical and empirical studies have shown that relative configurations of such channels are important to ecosystem stability. Our results suggest that functional groupings of plants based on photosynthetic pathway or other plant traits likely form the basis for food web compartments. By using diverse functional groups of plants for reclamation or restoration, practitioners may be able to aid the development of channels and thereby promote desired ecosystem states.     

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.     

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.     

C3 grasses have higher nutritional quality than C4 grasses under ambient and elevated atmospheric CO2

Grasses with the C₃ photosynthetic pathway are commonly considered to be more nutritious host plants than C₄ grasses, but the nutritional quality of C₃ grasses is also more greatly impacted by elevated atmospheric CO₂ than is that of C₄ grasses; C₃ grasses produce greater amounts of nonstructural carbohydrates and have greater declines in their nitrogen content than do C₄ grasses under elevated CO₂. Will C₃ grasses remain nutritionally superior to C₄ grasses under elevated CO₂ levels? We addressed this question by determining whether levels of protein in C₃ grasses decline to similar levels as in C₄ grasses, and whether total carbohydrate : protein ratios become similar in C₃ and C₄ grasses under elevated CO₂. In addition, we tested the hypothesis that, among the nonstructural carbohydrates in C₃ grasses, levels of fructan respond most strongly to elevated CO₂. Five C₃ and five C₄ grass species were grown from seed in outdoor open‐top chambers at ambient (370 ppm) or elevated (740 ppm) CO₂ for 2 months. As expected, a significant increase in sugars, starch and fructan in the C₃ grasses under elevated CO₂ was associated with a significant reduction in their protein levels, while protein levels in most C₄ grasses were little affected by elevated CO₂. However, this differential response of the two types of grasses was insufficient to reduce protein in C₃ grasses to the levels in C₄ grasses. Although levels of fructan in the C₃ grasses tripled under elevated CO₂, the amounts produced remained relatively low, both in absolute terms and as a fraction of the total nonstructural carbohydrates in the C₃ grasses. We conclude that C₃ grasses will generally remain more nutritious than C₄ grasses at elevated CO₂ concentrations, having higher levels of protein, nonstructural carbohydrates, and water, but lower levels of fiber and toughness, and lower total carbohydrate : protein ratios than C₄ grasses.     

The occurrence of chloroplast peripheral reticulum in grasses: a matter of phylogeny or a matter of function?

The chloroplast peripheral reticulum (PR) is a structure of unknown function. Some authors postulated that it is a characteristic feature of C₄ plants, although it was reported from C₃ species as well. It is unknown whether the occurrence of PR follows a phylogenetic (it is found in clades containing C₄ species, regardless of the photosynthetic type) or functional (photosynthetic pathway dependent) pattern. Here, we present a phylogenetically controlled analysis of the occurrence, form and functional aspects of PR in grasses. The occurrence of the PR follows a functional and not a phylogenetic pattern. Its most elaborated form (PR type I) is a unique feature of C₄ species. Although PR was found in some of the studied C₃ grasses, it was always less developed than PR in the chloroplasts of Kranz mesophyll cells of C₄ species. The size of PR in C₄ plants was found to increase when the plants were grown under low light intensity. Additional observations, such as a negative correlation between PR size and chloroplast surface and PR occurrence in vicinity of mitochondria or plasmodesmata, suggest that PR may play some role in C₄ metabolism.     

Differential herbivory on C3 versus C4 grasses by the grasshopper Ageneotettix deorum (Orthoptera: acrididae)

Summary The hypothesis that graminivorous grasshoppers select C₃ grasses over C₄ grasses was tested with experiments in the field. It was found that the generalist, graminivorous grasshopper Ageneotettix deorum typically chooses C₃ as opposed to C₄ grasses when both types are equally available. This preference is attributed to the differences in leaf anatomies of C₃ and C₄ plants since crude protein, water, lignin, fiber, and silicon content or the size of the individual leaves tested do not explain the observed feeding pattern. However, examination of the actual food plant selection of actual field inhabiting grasshoppers indicates that food plant prefernce may only be a minor component of food selection in natural settings.     

Distribution of C3 and C4 grasses at different altitudes in a temperate arid region of Argentina

Summary The distribution of native C₃ and C₄ grasses in a temperate arid region of Mendoza, Argentina, was studied in six areas at different altitudes. C₄ species predominate at low elevations in both relative species abundance and plant cover. At high elevations C₃ species are dominant in cover and composition. At medium altitudes (1100–1600 m) grass species composition is balanced but plant cover of C₃ species is greater. Of 31 genera in the whole area, 19 were C₄. Only the genera Stipa (C₃) and Aristida (C₄) were present in all the six areas surveyed. The pattern of grass distribution shows high correlation with evapotranspiration and temperature parameters, but low correlation with rainfall. The relation between grass distribution and different climatic parameters is discussed.     

Dynamic photo-inhibition and carbon gain in a C4 and a C3 grass native to high latitudes

C₄ plants are rare in the cool climates characteristic of high latitudes and altitudes, perhaps because of an enhanced susceptibility to photo‐inhibition at low temperatures relative to C₃ species. In the present study we tested the hypothesis that low‐temperature photo‐inhibition is more detrimental to carbon gain in the C₄ grass Muhlenbergia glomerata than the C₃ species Calamogrostis Canadensis. These grasses occur together in boreal fens in northern Canada. Plants were grown under cool (14/10 °C day/night) and warm (26/22 °C) temperatures before measurement of the light responses of photosynthesis and chlorophyll fluorescence at different temperatures. Cool growth temperatures led to reduced rates of photosynthesis in M. glomerata at all measurement temperatures, but had a smaller effect on the C₃ species. In both species the amount of xanthophyll cycle pigments increased when plants were grown at 14/10 °C, and in M. glomerata the xanthophyll epoxidation state was greatly reduced. The detrimental effect of low growth temperature on photosynthesis in M. glomerata was almost completely reversed by a 24‐h exposure to the warm‐temperature regime. These data indicate that reversible dynamic photo‐inhibition is a strategy by which C₄ species may tolerate cool climates and overcome the Rubisco limitation that is prevalent at low temperatures in C₄ plants.     

A comparative analysis of photosynthetic characteristics of hulless barley at two altitudes on the Tibetan Plateau

To determine the photosynthetic characteristics of C₃ plants and their sensitivity to CO₂ at different altitudes on the Tibetan Plateau, hulless barley (Hordeum vulgare L. ssp. vulgare) was grown at altitudes of 4,333 m and 3,688 m. Using gas-exchange measurements, photosynthetic parameters were simulated, including the maximum net photosynthesis (P _max) and the apparent quantum efficiency (α). Plants growing at higher altitude had higher net photosynthetic rates (P _N), photosynthesis parameters (P _max and α) and sensitivities to CO₂ enhancement than plants growing at lower altitude on the Tibetan Plateau. The enhancements of P _N, P _max, and α for plants growing at higher altitude, corresponding with 10 μmol(CO₂) mol⁻¹ increments, were approximately 0.20∼0.45%, 0.05∼0.20% and 0.12∼0.36% greater, respectively, than for plants growing at lower altitude, respectively, where CO₂ levels rose from 10 to 170 μmol(CO₂) mol⁻¹. Therefore, on the Tibetan Plateau, the changes in the photosynthetic capacities and the photosynthetic sensitivities to CO₂ observed in the C₃ plants grown above 3,688 m are likely to increase with altitude despite the decreasing CO₂ partial pressure.