Intraspecific variation in temperature dependence of gas exchange characteristics among Plantago asiatica ecotypes from different temperature regimes

There are large inter- and intraspecific differences in the temperature dependence of photosynthesis, but the physiological cause of the variation is poorly understood. Here, the temperature dependence of photosynthesis was examined in three ecotypes of Plantago asiatica transplanted from different latitudes, where the mean annual temperature varies between 7.5 and 16.8 degrees C. Plants were raised at 15 or 30 degrees C, and the CO(2) response of photosynthetic rates was determined at various temperatures. When plants were grown at 30 degrees C, no difference was found in the temperature dependence of photosynthesis among ecotypes. When plants were grown at 15 degrees C, ecotypes from a higher latitude maintained a relatively higher photosynthetic rate at low measurement temperatures. This difference was caused by a difference in the balance between the capacities of two processes, ribulose-1,5-bisphosphate regeneration (J(max)) and carboxylation (V(cmax)), which altered the limiting step of photosynthesis at low temperatures. The organization of photosynthetic proteins also varied among ecotypes. The ecotype from the highest latitude increased the J(max) : V(cmax) ratio with decreasing growth temperature, while that from the lowest latitude did not. It is concluded that nitrogen partitioning in the photosynthetic apparatus and its response to growth temperature were different among ecotypes, which caused an intraspecific variation in temperature dependence of photosynthesis.     

Seasonal differences in photosynthesis between the C3 and C4 subspecies of Alloteropsis semialata are offset by frost and drought

The regional abundance of C₄ grasses is strongly controlled by temperature, however, the role of precipitation is less clear. Progress in elucidating the direct effects of photosynthetic pathway on these climate relationships is hindered by the significant genetic divergence between major C₃ and C₄ grass lineages. We addressed this problem by examining seasonal climate responses of photosynthesis in Alloteropsis semialata , a unique grass species with both C₃ and C₄ subspecies. Experimental manipulation of rainfall in a common garden in South Africa tested the hypotheses that: (1) photosynthesis is greater in the C₄ than C₃ subspecies under high summer temperatures, but this pattern is reversed at low winter temperatures; and (2) the photosynthetic advantage of C₄ plants is enhanced during drought events. Measurements of leaf gas exchange over 2 years showed a significant photosynthetic advantage for the C₄ subspecies under irrigated conditions from spring through autumn. However, the C₄ leaves were killed by winter frost, while photosynthesis continued in the C₃ plants. Unexpectedly, the C₄ subspecies also lost its photosynthetic advantage during natural drought events, despite greater water-use efficiency under irrigated conditions. This study highlights previously unrecognized roles for climatic extremes in determining the ecological success of C₃ and C₄ grasses.     

Low-temperature photosynthetic performance of a C4 grass and a co-occurring C3 grass native to high latitudes

The photosynthetic performance of C₄ plants is generally inferior to that of C₃ species at low temperatures, but the reasons for this are unclear. The present study investigated the hypothesis that the capacity of Rubisco, which largely reflects Rubisco content, limits C₄ photosynthesis at suboptimal temperatures. Photosynthetic gas exchange, chlorophyll a fluorescence, and the in vitro activity of Rubisco between 5 and 35 °C were measured to examine the nature of the low‐temperature photosynthetic performance of the co‐occurring high latitude grasses, Muhlenbergia glomerata (C₄) and Calamogrostis canadensis (C₃). Plants were grown under cool (14/10 °C) and warm (26/22 °C) temperature regimes to examine whether acclimation to cool temperature alters patterns of photosynthetic limitation. Low‐temperature acclimation reduced photosynthetic rates in both species. The catalytic site concentration of Rubisco was approximately 5.0 and 20 µmol m^?2 in M. glomerata and C. canadensis, respectively, regardless of growth temperature. In both species, in vivo electron transport rates below the thermal optimum exceeded what was necessary to support photosynthesis. In warm‐grown C. canadensis, the photosynthesis rate below 15 °C was unaffected by a 90% reduction in O₂ content, indicating photosynthetic capacity was limited by the capacity of P_i‐regeneration. By contrast, the rate of photosynthesis in C. canadensis plants grown at the cooler temperatures was stimulated 20–30% by O₂ reduction, indicating the P_i‐regeneration limitation was removed during low‐temperature acclimation. In M. glomerata, in vitro Rubisco activity and gross CO₂ assimilation rate were equivalent below 25 °C, indicating that the capacity of the enzyme is a major rate limiting step during C₄ photosynthesis at cool temperatures.     

不同土壤水分条件下中国沙棘雌雄株光合作用、蒸腾作用及水分利用效率特征

采用LI-6400便携式光合系统对野外不同生境(沟底、坡面)和田间不同土壤水分条件下中国沙棘雌雄株的净光合速率和蒸腾速率及水分利用效率特征进行了观测,结果表明,中国沙棘雌雄株在水分条件较好的生境中均表现较强的生活力,净光合速率和蒸腾速率主要受光照强度和大气温度的影响,雄株表现出更高的光合、蒸腾、水分利用效率;在水分条件较差的生境中,雌雄株均通过降低蒸腾和提高水分利用效率来适应逆境,净光合速率和蒸腾速率的影响因子增多,雄株的光合速率大幅度下降,雌株仍保持较高的光合速率,雌株的水分利用效率高于雄株.可见,在反映植物瞬时生理变化的性状方面,雌株对土壤水分减少表现出了更强的适应性,雄株对土壤水分变化表现出了更大的敏感性.     

Interactive effects of elevated carbon dioxide and growth temperature on photosynthesis in cotton leaves

Cotton (Gossypium hirsutum L., cv DPL 5415) plants were grown in naturally lit environment chambers at day/night temperature regimes of 26/18 (T-26/18), 31/23 (T-31/23) and 36/28 °C (T-36/28) and CO2 concentrations of 350 (C-350), 450 (C-450) and 700 L L-1 (C-700). Net photosynthesis rates, stomatal conductance, transpiration, RuBP carboxylase activity and the foliar contents of starch and sucrose were measured during different growth stages. Net CO2 assimilation rates increased with increasing CO2 and temperature regimes. The enhancement of photosynthesis was from 24 mol CO2 m-2 s-1 (with C-350 and T-26/18) to 41 mol m-2 s-1 (with C-700 and T-36/28). Stomatal conductance decreased with increasing CO2 while it increased up to T-31/23 and then declined. The interactive effects of CO2 and temperature resulted in a 30% decrease in transpiration. Although the leaves grown in elevated CO2 had high starch and sucrose concentrations, their content decreased with increasing temperature. Increasing temperature from T-26/18 to 36/28 increased RuBP carboxylase activity in the order of 121, 172 and 190 mol mg-1 chl h-1 at C-350, C-450 and C-700 respectively. Our data suggest that leaf photosynthesis in cotton benefited more from CO_2 enrichment at warm temperatures than at low growth temperature regimes.     

高浓度二氧化碳对植物影响的研究进展

工业革命后全球大气CO2浓度持续上升,不仅对全球气候的变迁产生重大影响,而且对植物的形态、水分利用、蛋白质合成、光合、抗性、生长及生物量等都有不同程度的影响。高浓度CO2促进植物根、幼苗的生长,叶片增厚,降低气孔密度、气孔导度及蒸腾速率,增加水分利用效率、作物的产量及生物量,促进乙烯生物合成,增强植物的抗氧化能力。不同光合途径(C3、C4及CAM)及不同植被类型的植物对高浓度CO2的响应不同。长期和短期的高浓度CO2处理,植物响应方式有很大的差异,如短期高CO2处理使光合能力增强,而长期处理则使光合能力下调。     

Interactive Effects of Elevated CO2 and Growth Temperature on the Tolerance of Photosynthesis to Acute Heat Stress in C3 and C4 Species

Determining effects of elevated CO2 on the tolerance of photosynthesis to acute heat-stress (heat wave) is necessary for predicting plant responses to global warming, as photosynthesis is thermolabile and acute heat-stress and atmospheric CO2 will increase in the future. Few studies have examined this, and past results are variable, which may be due to methodological variation. To address this, we grew two C3 and two C4 species at current or elevated CO2 and three different growth temperatures (GT). We assessed photosynthetic thermotolerance in both unacclimated (basal tolerance) and preheat-stressed (preHS = acclimated) plants. In C3 species, basal thermotolerance of net photosynthesis (Pn) was increased In high CO2, but in C4 species, Pn thermotlerance was decreased by high CO2 (except Zea maya at low GT); CO2 effects in preHS plants were mostly small or absent, though high CO2 was detrimental in one C3 and one C4 species at warmer GT. Though high CO2 generally decreased stomatal conductance, decreases in Pn during heat stress were mostly due to non-stomatal effects. Photosystem II (PSII) efficiency was often decreased by high CO2 during heat stress, especially at high GT; CO2 effects on post-PSll electron transport were variable. Thus, high CO2 often affected photosynthetic theromotolerance, and the effects varied with photosynthetic pathway, growth temperature, and acclimation state. Most importantly, in heat-stressed plants at normal or warmer growth temperatures, high CO2 may often decrease, or not benefit as expected, tolerance of photosynthesis to acute heat stress. Therefore, interactive effects of elevated CO2 and warmer growth temperatures on acute heat tolerance may contribute to future changes in plant productivity, distribution, and diversity.     

Evolution of C4 plants: a new hypothesis for an interaction of CO2 and water relations mediated by plant hydraulics

C(4) photosynthesis has evolved more than 60 times as a carbon-concentrating mechanism to augment the ancestral C(3) photosynthetic pathway. The rate and the efficiency of photosynthesis are greater in the C(4) than C(3) type under atmospheric CO(2) depletion, high light and temperature, suggesting these factors as important selective agents. This hypothesis is consistent with comparative analyses of grasses, which indicate repeated evolutionary transitions from shaded forest to open habitats. However, such environmental transitions also impact strongly on plant-water relations. We hypothesize that excessive demand for water transport associated with low CO(2), high light and temperature would have selected for C(4) photosynthesis not only to increase the efficiency and rate of photosynthesis, but also as a water-conserving mechanism. Our proposal is supported by evidence from the literature and physiological models. The C(4) pathway allows high rates of photosynthesis at low stomatal conductance, even given low atmospheric CO(2). The resultant decrease in transpiration protects the hydraulic system, allowing stomata to remain open and photosynthesis to be sustained for longer under drying atmospheric and soil conditions. The evolution of C(4) photosynthesis therefore simultaneously improved plant carbon and water relations, conferring strong benefits as atmospheric CO(2) declined and ecological demand for water rose.     

内蒙古半干旱草原区沙地植物群落光合特征的动态研究

沙蒿(Artemisia intramongolica)群落是半干旱草原地区沙地的重要植被类型,分别在植物的生长前期、中期、盛期和后期采用便携式光合测定仪和大型同化分析仪测定了沙蒿叶片和沙蒿群落的光合动态。单叶和群落的光合速率日进程类型随气候的不同而异,瞬时光合速率主要决定于光合有效辐射强度(PAR)。土壤干旱大大降低了单叶和群落的光合能力,晴天土壤湿润时气温和空气湿度控制着叶片的光合速率,午间大气湿度降低是光合午休的主要外因。叶片的蒸腾速率与气温呈显著线性相关,植物的光能和水分利用效率也主要取决于PAR和气温,随着PAR和气温的升高利用效率下降。沙蒿叶片光能利用效率在后期也能保持较高水平。沙蒿对土壤干旱和高温具有一定的适应性,在土壤湿润时能迅速提高光合速率,形成较大的生物量。但是沙蒿的蒸腾速率高,水分利用效率低。研究认为,沙蒿通过对土壤干旱和高温的忍耐机制而保持长时间较高的光能利用效率,并在土壤湿润时迅速提高光合能力和积累干物质来适应半干旱的沙地环境,而且依靠高蒸腾速率和强的水分吸收能力来竞争性抑制其他植物的生长。     

Photosynthetic responses of two heliotropic legumes from contrasting habitats

Abstract Photosynthetic responses to light, temperature and leaf-to-air water vapour concentration deficit for Strophostyles helvola (L.) Ell. in an open beach site, and Amphicarpa bracteata (L.) Ell. in two deciduous forest sites were quantified. Photosaturated rates of net CO₂ assimilation were 52.1 ± 4.6, 11.0 ± 1.6 and 4.1 ± 0.3 μmol m^?2s^?2 for plants in beach, roadside and closed canopy sites, respectively. In terms of photosynthesis, plants in the beach site were more tolerant of higher leaf temperatures and water vapour concentration deficits than were plants in forested sites. Heliotropic leaf movements in the beach site reduced calculated total daily transpiration by 2%, increased total daily carbon gain by 8% and reduced the transpiration ratio by 9% relative to an horizontal leaf. During long-term sunflecks in forested sites, heliotropism reduced transpiration by 15%, increased carbon gain by 71% and reduced the transpiration ratio by 50% relative to an horizontal leaf. We hypothesize that heliotropic leaf movements in mesic, high-light, low-nitrogen habitats may increase carbon return on nitrogen investment in photosynthesis, while heliotropism in canopy gaps may represent a morphological mechanism to reduce damage to low-light acclimated photosynthetic systems during long-term sunflecks.     

Diffusive and metabolic limitations to photosynthesis under drought and salinity in C(3) plants

Drought and salinity are two widespread environmental conditions leading to low water availability for plants. Low water availability is considered the main environmental factor limiting photosynthesis and, consequently, plant growth and yield worldwide. There has been a long-standing controversy as to whether drought and salt stresses mainly limit photosynthesis through diffusive resistances or by metabolic impairment. Reviewing in vitro and in vivo measurements, it is concluded that salt and drought stress predominantly affect diffusion of CO(2) in the leaves through a decrease of stomatal and mesophyll conductances, but not the biochemical capacity to assimilate CO(2), at mild to rather severe stress levels. The general failure of metabolism observed at more severe stress suggests the occurrence of secondary oxidative stresses, particularly under high-light conditions. Estimates of photosynthetic limitations based on the photosynthetic response to intercellular CO(2) may lead to artefactual conclusions, even if patchy stomatal closure and the relative increase of cuticular conductance are taken into account, as decreasing mesophyll conductance can cause the CO(2) concentration in chloroplasts of stressed leaves to be considerably lower than the intercellular CO(2) concentration. Measurements based on the photosynthetic response to chloroplast CO(2) often confirm that the photosynthetic capacity is preserved but photosynthesis is limited by diffusive resistances in drought and salt-stressed leaves.     

植物光合作用对大气CO2浓度升高的反应

近年来大气中ＣＯ２浓度急剧增加使人们重新对研究ＣＯ２浓度升高对植物光合作用影响感兴趣。预计在未来的１００ａ中,大气ＣＯ２浓度还将不断增长并达到当今的２倍。ＣＯ２排放量的增加不仅加剧了地球上的温室效应,也将改变全球生态系统中碳的平衡。离浓度ＣＯ２对植物光剑作用的影响表现为短期和长期效应。短时间地供给高浓度ＣＯ２促进阿 光合作用,而长时间生长在高浓度ＣＯ２下抒使某些植物光合能力下降,出现了光合适应现象     

The sensitivity of C3 photosynthesis to increasing CO2 concentration: a theoretical analysis of its dependence on temperature and background CO2 concentration

The atmospheric CO₂ concentration has increased from the pre-industrial concentration of about 280 μmol mol⁻¹ to its present concentration of over 350 μmol mol⁻¹, and continues to increase. As the rate of photosynthesis in C₃ plants is strongly dependent on CO₂ concentration, this should have a marked effect on photosynthesis, and hence on plant growth and productivity. The magnitude of photo-synthetic responses can be calculated based on the well-developed theory of photosynthetic response to intercellular CO₂ concentration. A simple biochemically based model of photosynthesis was coupled to a model of stomatal conductance to calculate photosynthetic responses to ambient CO₂ concentration. In the combined model, photosynthesis was much more responsive to CO₂ at high than at low temperatures. At 350 μmol mol⁻¹, photosynthesis at 35°C reached 51% of the rate that would have been possible with non-limiting CO₂, whereas at 5°C, 77% of the CO₂ non-limited rate was attained. Relative CO₂ sensitivity also became smaller at elevated CO₂, as CO₂ concentration increased towards saturation. As photosynthesis was far from being saturated at the current ambient CO₂ concentration, considerable further gains in photosynthesis were predicted through continuing increases in CO₂ concentration. The strong interaction with temperature also leads to photosynthesis in different global regions experiencing very different sensitivities to increasing CO₂ concentrations.     

Effects of nitrogen supply and high temperature on the growth and physiology of the chickpea

Chickpeas were grown with or without nitrate nitrogen feeding, or nodulated with Rhizobium leguminosarum. High [40°C day, 25°C night (HT)] and moderate [25°C day, 177°C night (LT)] temperature regimes were employed during growth. Growth rates, photosynthetic capacity and enzymes of carbon and nitrogen metabolism were monitored to assess the acclimatory capacity of the chickpea. Initial growth rates were stimulated by high temperatures, particularly in nitrate-fed and nodulated plants. Older HT plants had fewer laterals, smaller leaves, and fewer flowers were produced than in LT plants. There was some indication of an acclimation of photosynthesis to high temperatures and this was independent of nitrogen supply. Rubisco activity was increased by high growth temperatures. However, HT plants also had higher transpiration rates and lower water use efficiency than LT plants both in respective growth conditions and when compared in a common condition. High temperatures reduced shoot nitrate reductase activity but had little effect on root activity, which was the same if not greater than activity in LT roots. The amino acid, asparagine, was found at high concentrations in all treatments. Concentrations were maintained throughout growth in HT plants but declined with age in LT plants.     

山西太岳山辽东栎的光合特性

对自然生长的辽东栎林木用 LI-640 0测定其光合作用 ,实验包括以下内容 :在完全接近自然生长的条件下测定辽东栎叶片光合作用的日变化规律 ;控制光合有效辐射强度 ,测定叶片在不同 CO2 浓度下光合作用的变化规律 ;控制温度比对应时间段的温度高 2～ 4℃ ,并控制 CO2 浓度在 3 75～ 70 0 ml· m- 3间变化 ,测定辽东栎光合作用的变化情况。结果如下 :1自然状态下辽东栎光合作用的日变化有两个光合速率峰 ,峰高接近 ,上午的光合速率峰持续时间较下午的长。叶片气孔导度与光合速率间有很强的正相关 ,且对胞内 CO2 浓度和蒸腾速率有较大影响。气孔导度与叶片水压亏缺呈相反的变化趋势 ,而水压亏缺受光合有效辐射、气温的影响较大。 2在控制片温度、光合有效辐射不变的情况下 ,辽东栎的叶片水压亏缺变化幅度很小 ,气孔导度变化主要受 CO2 浓度的调节 ,表现出在目前 CO2 浓度至加倍 CO2 浓度下 ,随浓度增加而增大的趋势 ,与之对应光合速率增大 ,同时光补偿点也有所提高。当 CO2 浓度超过 80 0 ml· m- 3继续上升时 ,气孔导度、光合速率均下降 ,光补偿点继续升高。3对辽东栎叶片进行升温和高 CO2 浓度处理 ,在试验进行时的温度 (2 8～ 3 7℃ )下 ,发现在 3 75～ 70 0 ml· m- 3的 CO2 浓度范围内 ,升温均导致光     

The temperature response of C(3) and C(4) photosynthesis

We review the current understanding of the temperature responses of C(3) and C(4) photosynthesis across thermal ranges that do not harm the photosynthetic apparatus. In C(3) species, photosynthesis is classically considered to be limited by the capacities of ribulose 1.5-bisphosphate carboxylase/oxygenase (Rubisco), ribulose bisphosphate (RuBP) regeneration or P(i) regeneration. Using both theoretical and empirical evidence, we describe the temperature response of instantaneous net CO(2) assimilation rate (A) in terms of these limitations, and evaluate possible limitations on A at elevated temperatures arising from heat-induced lability of Rubisco activase. In C(3) plants, Rubisco capacity is the predominant limitation on A across a wide range of temperatures at low CO(2) (<300 microbar), while at elevated CO(2), the limitation shifts to P(i) regeneration capacity at suboptimal temperatures, and either electron transport capacity or Rubisco activase capacity at supraoptimal temperatures. In C(4) plants, Rubisco capacity limits A below 20 degrees C in chilling-tolerant species, but the control over A at elevated temperature remains uncertain. Acclimation of C(3) photosynthesis to suboptimal growth temperature is commonly associated with a disproportional enhancement of the P(i) regeneration capacity. Above the thermal optimum, acclimation of A to increasing growth temperature is associated with increased electron transport capacity and/or greater heat stability of Rubisco activase. In many C(4) species from warm habitats, acclimation to cooler growth conditions increases levels of Rubisco and C(4) cycle enzymes which then enhance A below the thermal optimum. By contrast, few C(4) species adapted to cooler habitats increase Rubisco content during acclimation to reduced growth temperature; as a result, A changes little at suboptimal temperatures. Global change is likely to cause a widespread shift in patterns of photosynthetic limitation in higher plants. Limitations in electron transport and Rubisco activase capacity should be more common in the warmer, high CO(2) conditions expected by the end of the century.     

松嫩草地全叶马兰夏季与秋季光合及蒸腾作用的比较

在生长季晴天条件下,夏季和秋季松嫩草地全叶马兰的光合,蒸腾作用的日变化均为双峰曲线,但不同季节有所差异,夏季日均净光合速率与蒸腾速率均高于秋季,光合和蒸腾作用与环境因子的植物内部因子之间有密切关系。分析表明,叶片净光合速率与有效光辐射呈极显著相关,与气孔阻力,胞间CO2呈负相关;蒸腾速率与有效光辐射呈极显著相关,与叶温,饱和差呈正相关,与气孔阻力,胞间CO2浓度呈负相关,有效光辐射是影响光合和蒸腾作用诸因子中的主导因子,而气孔阻力变化则在调节光合和蒸腾中起着重要作用,蒸腾速率午降主要由于光辐射强,叶温高,湿度低,植物体缺水,气孔部分关闭所致。     

Photosynthesis and transpiration by young Larix kaempferi trees: C3 responses to light and temperature

The effects of photon flux density and temperature on net photosynthesis and transpiration rates of mature and immature leaves of three-year-old Japanese larch Larix kaempferi (Lamb.) Sarg. trees were determined with an infrared, differential open gas analysis system. Net photosynthetic response to increasing photon flux densities was similar for different foliage positions and stage of maturity. Light compensation was between 25 and 50 μmol m⁻² s⁻¹. Rates of photosynthesis increased rapidly at photon flux densities above the compensation level and became saturated between 800 and 1000 μmol m⁻² s⁻¹. Transpiration rates at constant temperature likewise increased with increasing photon flux density, and leveled off between 800 and 1000 μmol m⁻² s⁻¹. Photosynthetic response to temperature was determined in saturating light and was similar for all foliage positions; it increased steadily from low temperatures to an optimum range betweeen 15 and 21°C and then decreased rapidly above 21°C. Transpiration rate, however, increased continuously with rising temperature up to the experimental maximum. CO₂ compensation concentrations for mature foliage varied between 58 and 59 μl l⁻¹; however, foliage borne at the apex of the terminal leader compensated at 75 μl l⁻¹. None of these data support the claim that Japanese larch possesses C₄ photosynthetic characteristics.     

Photosynthesis and cannabinoid content of temperate and tropical populations of Cannabis sativa

Four populations of Cannabis sativa L. grown from seeds collected in Panama, Jamaica, Nepal, and east central Illinois were grown under controlled conditions in growth chambers. One set was grown under warm conditions (32° day and 23° night) and the other set was grown under lower temperatures (23° day and 16° night). CO₂ exchange and transpiration were examined under various temperatures and light intensities. Observations on growth, and analyses for chlorophyll and Δ₁THC (tetrahydrocannabinol) content were made. Under warm growth conditions, the central Illinois population had the highest photosynthetic rate at all temperatures investigated. The Nepal population had intermediate rates, while the Jamaica and the Panama populations had the lowest rate. The Jamaica and Panama populations had insignificant changes in photosynthetic response to changes in temperatures between 15° and 30°. Under cool growing conditions the central Illinois population had the highest rate of photosynthesis with a definite peak at 25°. Nepal plants had intermediate rates of photosynthesis, while the Panama and Jamaica populations had the lowest rate. Differences in chlorophyll and drug content were also significant between these populations. From these data it is suggested that the four populations can be grouped into different ecotypes genetically adapted to their respective environments.     

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.