Invasive grasses of sub-Antarctic Marion Island respond to increasing temperatures at the expense of chilling tolerance.

BACKGROUND AND AIMS: Global warming has large effects on the performance and spatial distribution of plants, and increasingly facilitates the spread of invasive species. Particularly vulnerable is the vegetation of cold environments where indigenous plants selected for cold tolerance can have reduced phenotypic plasticity and associated lower capacity to respond to warming temperatures. In contrast, invasive species can be phenotypically plastic and respond positively to climate change, but at the expense of stress tolerance. METHODS: We investigate this trade-off in traits, measuring the photosynthetic response to warming, chilling tolerance and specific leaf area (SLA) of Pooid grasses. We compare this between invasive and non-invasive grasses and correlate this to their range expansions on a cold sub-Antarctic island that has warmed significantly in the past five decades. We determined whether these responses remained consistent after temperature acclimation. KEY RESULTS: Invasive species responded strongly to warming, increasing photosynthetic rates by up to 2-fold, while non-invasive species did not respond. The response was associated with increased stomatal conductance, but not with modified photosynthetic metabolism. Electrolyte leakage and SLA were higher in invasive than in non-invasive species. Acclimation altered the photosynthetic response and invasive species responded to warm temperatures irrespective of acclimation, while non-invasive species responded only after acclimation to warm temperature. CONCLUSIONS: Traits scaled linearly with rates of range expansion and demonstrate that under sub-Antarctic conditions, anthropogenic warming over the last 50 years may have favoured species with greater capacity to respond photosynthetically to warming to the detriment of species that cannot, and negated the advantage that chilling tolerance would have conferred on endemic species in the past. This suggests that species of cold ecosystems could be particularly vulnerable to warming as selection for stress tolerance has limited their responsiveness to environmental change, while introduced invasive species may have no such limitations. We show mechanistic evidence of the physiology that underpins an apparent trade-off between warming and chilling tolerance traits.

Frequent fires prime plant developmental responses to burning.

Coping with temporal variation in fire requires plants to have plasticity in traits that promote persistence, but how plastic responses to current conditions are affected by past fire exposure remains unknown. We investigate phenotypic divergence between populations of four resprouting grasses exposed to differing experimental fire regimes (annually burnt or unburnt for greater than 35 years) and test whether divergence persists after plants are grown in a common environment for 1 year. Traits relating to flowering and biomass allocation were measured before plants were experimentally burnt, and their regrowth was tracked. Genetic differentiation between populations was investigated for a subset of individuals. Historic fire frequency influenced traits relating to flowering and below-ground investment. Previously burnt plants produced more inflorescences and invested proportionally more biomass below ground, suggesting a greater capacity for recruitment and resprouting than unburnt individuals. Tiller-scale regrowth rate did not differ between treatments, but prior fire exposure enhanced total regrown biomass in two species. We found no consistent genetic differences between populations suggesting trait differences arose from developmental plasticity. Grass development is influenced by prior fire exposure, independent of current environmental conditions. This priming response to fire, resulting in adaptive trait changes, may produce communities more resistant to future fire regime changes.

C4 anatomy can evolve via a single developmental change.

C4 photosynthesis is a complex trait that boosts productivity in warm environments. Paradoxically, it evolved independently in numerous plant lineages, despite requiring specialised leaf anatomy. The anatomical modifications underlying C4 evolution have previously been evaluated through interspecific comparisons, which capture numerous changes besides those needed for C4 functionality. Here, we quantify the anatomical changes accompanying the transition between non-C4 and C4 phenotypes by sampling widely across the continuum of leaf anatomical traits in the grass Alloteropsis semialata. Within this species, the only trait that is shared among and specific to C4 individuals is an increase in vein density, driven specifically by minor vein development that yields multiple secondary effects facilitating C4 function. For species with the necessary anatomical preconditions, developmental proliferation of veins can therefore be sufficient to produce a functional C4 leaf anatomy, creating an evolutionary entry point to complex C4 syndromes that can become more specialised.

Preference for C4 shade grasses increases hatchling performance in the butterfly, Bicyclus safitza.

The Miocene radiation of C4 grasses under high-temperature and low ambient CO 2 levels occurred alongside the transformation of a largely forested landscape into savanna. This inevitably changed the host plant regime of herbivores, and the simultaneous diversification of many consumer lineages, including Bicyclus butterflies in Africa, suggests that the radiations of grasses and grazers may be evolutionary linked. We examined mechanisms for this plant-herbivore interaction with the grass-feeding Bicyclus safitza in South Africa. In a controlled environment, we tested oviposition preference and hatchling performance on local grasses with C3 or C4 photosynthetic pathways that grow either in open or shaded habitats. We predicted preference for C3 plants due to a hypothesized lower processing cost and higher palatability to herbivores. In contrast, we found that females preferred C4 shade grasses rather than either C4 grasses from open habitats or C3 grasses. The oviposition preference broadly followed hatchling performance, although hatchling survival was equally good on C4 or C3 shade grasses. This finding was explained by leaf toughness; shade grasses were softer than grasses from open habitats. Field monitoring revealed a preference of adults for shaded habitats, and stable isotope analysis of field-sampled individuals confirmed their preference for C4 grasses as host plants. Our findings suggest that plant-herbivore interactions can influence the direction of selection in a grass-feeding butterfly. Based on this work, we postulate future research to test whether these interactions more generally contribute to radiations in herbivorous insects via expansions into new, unexploited ecological niches.

Determinants of flammability in savanna grass species.

Tropical grasses fuel the majority of fires on Earth. In fire-prone landscapes, enhanced flammability may be adaptive for grasses via the maintenance of an open canopy and an increase in spatiotemporal opportunities for recruitment and regeneration. In addition, by burning intensely but briefly, high flammability may protect resprouting buds from lethal temperatures. Despite these potential benefits of high flammability to fire-prone grasses, variation in flammability among grass species, and how trait differences underpin this variation, remains unknown.By burning leaves and plant parts, we experimentally determined how five plant traits (biomass quantity, biomass density, biomass moisture content, leaf surface-area-to-volume ratio and leaf effective heat of combustion) combined to determine the three components of flammability (ignitability, sustainability and combustibility) at the leaf and plant scales in 25 grass species of fire-prone South African grasslands at a time of peak fire occurrence. The influence of evolutionary history on flammability was assessed based on a phylogeny built here for the study species.Grass species differed significantly in all components of flammability. Accounting for evolutionary history helped to explain patterns in leaf-scale combustibility and sustainability. The five measured plant traits predicted components of flammability, particularly leaf ignitability and plant combustibility in which 70% and 58% of variation, respectively, could be explained by a combination of the traits. Total above-ground biomass was a key driver of combustibility and sustainability with high biomass species burning more intensely and for longer, and producing the highest predicted fire spread rates. Moisture content was the main influence on ignitability, where species with higher moisture contents took longer to ignite and once alight burnt at a slower rate. Biomass density, leaf surface-area-to-volume ratio and leaf effective heat of combustion were weaker predictors of flammability components. Synthesis. We demonstrate that grass flammability is predicted from easily measurable plant functional traits and is influenced by evolutionary history with some components showing phylogenetic signal. Grasses are not homogenous fuels to fire. Rather, species differ in functional traits that in turn demonstrably influence flammability. This diversity is consistent with the idea that flammability may be an adaptive trait for grasses of fire-prone ecosystems.

Evolutionary implications of C3 -C4 intermediates in the grass Alloteropsis semialata.

C4 photosynthesis is a complex trait resulting from a series of anatomical and biochemical modifications to the ancestral C3 pathway. It is thought to evolve in a stepwise manner, creating intermediates with different combinations of C4 -like components. Determining the adaptive value of these components is key to understanding how C4 photosynthesis can gradually assemble through natural selection. Here, we decompose the photosynthetic phenotypes of numerous individuals of the grass Alloteropsis semialata, the only species known to include both C3 and C4 genotypes. Analyses of δ(13) C, physiology and leaf anatomy demonstrate for the first time the existence of physiological C3 -C4 intermediate individuals in the species. Based on previous phylogenetic analyses, the C3 -C4 individuals are not hybrids between the C3 and C4 genotypes analysed, but instead belong to a distinct genetic lineage, and might have given rise to C4 descendants. C3 A. semialata, present in colder climates, likely represents a reversal from a C3 -C4 intermediate state, indicating that, unlike C4 photosynthesis, evolution of the C3 -C4 phenotype is not irreversible.

Photosynthetic innovation broadens the niche within a single species.

Adaptation to changing environments often requires novel traits, but how such traits directly affect the ecological niche remains poorly understood. Multiple plant lineages have evolved C4 photosynthesis, a combination of anatomical and biochemical novelties predicted to increase productivity in warm and arid conditions. Here, we infer the dispersal history across geographical and environmental space in the only known species with both C4 and non-C4 genotypes, the grass Alloteropsis semialata. While non-C4 individuals remained confined to a limited geographic area and restricted ecological conditions, C4 individuals dispersed across three continents and into an expanded range of environments, encompassing the ancestral one. This first intraspecific investigation of C4 evolutionary ecology shows that, in otherwise similar plants, C4 photosynthesis does not shift the ecological niche, but broadens it, allowing dispersal into diverse conditions and over long distances. Over macroevolutionary timescales, this immediate effect can be blurred by subsequent specialisation towards more extreme niches.

Physiological advantages of C4 grasses in the field: a comparative experiment demonstrating the importance of drought.

Global climate change is expected to shift regional rainfall patterns, influencing species distributions where they depend on water availability. Comparative studies have demonstrated that C4 grasses inhabit drier habitats than C3 relatives, but that both C3 and C4 photosynthesis are susceptible to drought. However, C4 plants may show advantages in hydraulic performance in dry environments. We investigated the effects of seasonal variation in water availability on leaf physiology, using a common garden experiment in the Eastern Cape of South Africa to compare 12 locally occurring grass species from C4 and C3 sister lineages. Photosynthesis was always higher in the C4 than C3 grasses across every month, but the difference was not statistically significant during the wettest months. Surprisingly, stomatal conductance was typically lower in the C3 than C4 grasses, with the peak monthly average for C3 species being similar to that of C4 leaves. In water-limited, rain-fed plots, the photosynthesis of C4 leaves was between 2.0 and 7.4 µmol m(-2) s(-1) higher, stomatal conductance almost double, and transpiration 60% higher than for C3 plants. Although C4 average instantaneous water-use efficiencies were higher (2.4-8.1 mmol mol(-1)) than C3 averages (0.7-6.8 mmol mol(-1)), differences were not as great as we expected and were statistically significant only as drought became established. Photosynthesis declined earlier during drought among C3 than C4 species, coincident with decreases in stomatal conductance and transpiration. Eventual decreases in photosynthesis among C4 plants were linked with declining midday leaf water potentials. However, during the same phase of drought, C3 species showed significant decreases in hydrodynamic gradients that suggested hydraulic failure. Thus, our results indicate that stomatal and hydraulic behaviour during drought enhances the differences in photosynthesis between C4 and C3 species. We suggest that these drought responses are important for understanding the advantages of C4 photosynthesis under field conditions.

How succulent leaves of Aizoaceae avoid mesophyll conductance limitations of photosynthesis and survive drought.

In several taxa, increasing leaf succulence has been associated with decreasing mesophyll conductance (g M) and an increasing dependence on Crassulacean acid metabolism (CAM). However, in succulent Aizoaceae, the photosynthetic tissues are adjacent to the leaf surfaces with an internal achlorophyllous hydrenchyma. It was hypothesized that this arrangement increases g M, obviating a strong dependence on CAM, while the hydrenchyma stores water and nutrients, both of which would only be sporadically available in highly episodic environments. These predictions were tested with species from the Aizoaceae with a 5-fold variation in leaf succulence. It was shown that g M values, derived from the response of photosynthesis to intercellular CO2 concentration (A:C i), were independent of succulence, and that foliar photosynthate δ(13)C values were typical of C3, but not CAM photosynthesis. Under water stress, the degree of leaf succulence was positively correlated with an increasing ability to buffer photosynthetic capacity over several hours and to maintain light reaction integrity over several days. This was associated with decreased rates of water loss, rather than tolerance of lower leaf water contents. Additionally, the hydrenchyma contained ~26% of the leaf nitrogen content, possibly providing a nutrient reservoir. Thus the intermittent use of C3 photosynthesis interspersed with periods of no positive carbon assimilation is an alternative strategy to CAM for succulent taxa (Crassulaceae and Aizoaceae) which occur sympatrically in the Cape Floristic Region of South Africa.

Photosynthetic acclimation and resource use by the C3 and C4 subspecies of Alloteropsis semialata in low CO2 atmospheres.

During the past 25 Myr, partial pressures of atmospheric CO2 (Ca ) imposed a greater limitation on C3 than C4 photosynthesis. This could have important downstream consequences for plant nitrogen economy and biomass allocation. Here, we report the first phylogenetically controlled comparison of the integrated effects of subambient Ca on photosynthesis, growth and nitrogen allocation patterns, comparing the C3 and C4 subspecies of Alloteropsis semialata. Plant size decreased more in the C3 than C4 subspecies at low Ca , but nitrogen pool sizes were unchanged, and nitrogen concentrations increased across all plant partitions. The C3, but not C4 subspecies, preferentially allocated biomass to leaves and increased specific leaf area at low Ca . In the C3 subspecies, increased leaf nitrogen was linked to photosynthetic acclimation at the interglacial Ca , mediated via higher photosynthetic capacity combined with greater stomatal conductance. Glacial Ca further increased the biochemical acclimation and nitrogen concentrations in the C3 subspecies, but these were insufficient to maintain photosynthetic rates. In contrast, the C4 subspecies maintained photosynthetic rates, nitrogen- and water-use efficiencies and plant biomass at interglacial and glacial Ca with minimal physiological adjustment. At low Ca , the C4 carbon-concentrating mechanism therefore offered a significant advantage over the C3 type for carbon acquisition at the whole-plant scale, apparently mediated via nitrogen economy and water loss. A limiting nutrient supply damped the biomass responses to Ca and increased the C4 advantage across all Ca treatments. Findings highlight the importance of considering leaf responses in the context of the whole plant, and show that carbon limitation may be offset at the expense of greater plant demand for soil resources such as nitrogen and water. Results show that the combined effects of low CO2 and resource limitation benefit C4 plants over C3 plants in glacial-interglacial environments, but that this advantage is lessened under anthropogenic conditions.

Fire and fire-adapted vegetation promoted C4 expansion in the late Miocene.

Large proportions of the Earth's land surface are covered by biomes dominated by C(4) grasses. These C(4)-dominated biomes originated during the late Miocene, 3-8 million years ago (Ma), but there is evidence that C(4) grasses evolved some 20 Ma earlier during the early Miocene/Oligocene. Explanations for this lag between evolution and expansion invoke changes in atmospheric CO(2), seasonality of climate and fire. However, there is still no consensus about which of these factors triggered C(4) grassland expansion. We use a vegetation model, the adaptive dynamic global vegetation model (aDGVM), to test how CO(2), temperature, precipitation, fire and the tolerance of vegetation to fire influence C(4) grassland expansion. Simulations are forced with late Miocene climates generated with the Hadley Centre coupled ocean-atmosphere-vegetation general circulation model. We show that physiological differences between the C(3) and C(4) photosynthetic pathways cannot explain C(4) grass invasion into forests, but that fire is a crucial driver. Fire-promoting plant traits serve to expand the climate space in which C(4)-dominated biomes can persist. We propose that three mechanisms were involved in C(4) expansion: the physiological advantage of C(4) grasses under low atmospheric CO(2) allowed them to invade C(3) grasslands; fire allowed grasses to invade forests; and the evolution of fire-resistant savanna trees expanded the climate space that savannas can invade.

Ecophysiological traits in C3 and C4 grasses: a phylogenetically controlled screening experiment.

Experimental evidence demonstrates a higher efficiency of water and nitrogen use in C(4) compared with C(3) plants, which is hypothesized to drive differences in biomass allocation between C(3) and C(4) species. However, recent work shows that contrasts between C(3) and C(4) grasses may be misinterpreted without phylogenetic control. Here, we compared leaf physiology and growth in multiple lineages of C(3) and C(4) grasses sampled from a monophyletic clade, and asked the following question: which ecophysiological traits differ consistently between photosynthetic types, and which vary among lineages? C(4) species had lower stomatal conductance and water potential deficits, and higher water-use efficiency than C(3) species. Photosynthesis and nitrogen-use efficiency were also greater in C(4) species, varying markedly between clades. Contrary to previous studies, leaf nitrogen concentration was similar in C(4) and C(3) types. Canopy mass and area were greater, and root mass smaller, in the tribe Paniceae than in most other lineages. The size of this phylogenetic effect on biomass partitioning was greater in the C(4) NADP-me species than in species of other types. Our results show that the phylogenetic diversity underlying C(4) photosynthesis is critical to understanding its functional consequences. Phylogenetic bias is therefore a crucial factor to be considered when comparing the ecophysiology of C(3) and C(4) species.

A molecular phylogeny of the genus Alloteropsis (Panicoideae, Poaceae) suggests an evolutionary reversion from C4 to C3 photosynthesis.

BACKGROUND AND AIMS: The grass Alloteropsis semialata is the only plant species with both C(3) and C(4) subspecies. It therefore offers excellent potential as a model system for investigating the genetics, physiology and ecological significance of the C(4) photosynthetic pathway. Here, a molecular phylogeny of the genus Alloteropsis is constructed to: (a) confirm the close relationship between the C(3) and C(4) subspecies of A. semialata; and (b) infer evolutionary relationships between species within the Alloteropsis genus. METHODS: The chloroplast gene ndhF was sequenced from 12 individuals, representing both subspecies of A. semialata and all four of the other species in the genus. ndhF sequences were added to those previously sequenced from the Panicoideae, and used to construct a phylogenetic tree. KEY RESULTS: The phylogeny confirms that the two subspecies of A. semialata are among the most recently diverging lineages of C(3) and C(4) taxa currently recognized within the Panicoideae. Furthermore, the position of the C(3) subspecies of A. semialata within the Alloteropsis genus is consistent with the hypothesis that its physiology represents a reversion from C(4) photosynthesis. The data point to a similar evolutionary event in the Panicum stenodes-P. caricoides-P. mertensii clade. The Alloteropsis genus is monophyletic and occurs in a clade with remarkable diversity of photosynthetic biochemistry and leaf anatomy. CONCLUSIONS: These results confirm the utility of A. semialata as a model system for investigating C(3) and C(4) physiology, and provide molecular data that are consistent with reversions from C(4) to C(3) photosynthesis in two separate clades. It is suggested that further phylogenetic and functional investigations of the Alloteropsis genus and closely related taxa are likely to shed new light on the mechanisms and intermediate stages underlying photosynthetic pathway evolution.

Biomass reallocation and the mobilization of leaf resources support dune plant growth after sand burial.

The stimulation of dune plant growth in response to burial is a vital attribute allowing survival in areas of mobile sand. Numerous resource-related and physiological mechanisms of growth stimulation have been suggested in the past, but few have been tested comparatively. Manipulation experiments using Scaevola plumieri, an important subtropical coastal dune forming species, demonstrated that physiological shifts were of great importance in determining the nature of the stimulation response to burial. The production of stem length and replacement of leaf area were stimulated by burial, whereas net mass production was similar between buried and unburied treatments. Remobilization of buried leaf resources, seasonal effects, and a shift in biomass allocation to stem production played the greatest role in the compensatory growth response. Other factors, such as increased soil nutrients, changes in photosynthesis, and changes in the costs of producing tissue were of less importance. Thus, the stimulated growth of species adapted to live on mobile dunes is explained by a number of resource-related and physiological mechanisms acting in concert.

Low temperature effects on leaf physiology and survivorship in the C3 and C4 subspecies of Alloteropsis semialata.

The species richness of C(4) grasses is strongly correlated with temperature, with C(4) species dominating subtropical ecosystems and C(3) types predominating in cooler climates. Here, the effects of low temperatures on C(4) and C(3) grasses are compared, controlling for phylogenetic effects by using Alloteropsis semialata, a unique species with C(4) and C(3) subspecies. Controlled environment and common garden experiments tested the hypotheses that: (i) photosynthesis and growth are greater in the C(4) than the C(3) subspecies at high temperatures, but this advantage is reversed below 20 degrees C; and (ii) chilling-induced photoinhibition and light-mediated freezing injury of leaves occur at higher temperature thresholds in the C(4) than the C(3) plants. Measurements of leaf growth and photosynthesis showed the expected advantages of the C(4) pathway over the C(3) type at high temperatures. These declined with temperature, but were not completely lost until 15 degrees C, and there was no evidence of a reversal to give a C(3) advantage. Chronic chilling (5-15 degrees C) or acute freezing events induced a comparable degree of photodamage in illuminated leaves of both subspecies. Similarly, freezing caused high rates of mortality in the unhardened leaves of both subtypes. However, a 2-week chilling treatment prior to these freezing events halved injury in the C(3) but not the C(4) subspecies, suggesting that C(4) leaves lacked the capacity for cold acclimation. These results therefore suggest that C(3) members of this subtropical species may gain an advantage over their C(4) counterparts at low temperatures via protection from freezing injury rather than higher photosynthetic rates.

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

The regional abundance of C(4) 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 C(3) and C(4) grass lineages. We addressed this problem by examining seasonal climate responses of photosynthesis in Alloteropsis semialata, a unique grass species with both C(3) and C(4) subspecies. Experimental manipulation of rainfall in a common garden in South Africa tested the hypotheses that: (1) photosynthesis is greater in the C(4) than C(3) subspecies under high summer temperatures, but this pattern is reversed at low winter temperatures; and (2) the photosynthetic advantage of C(4) plants is enhanced during drought events. Measurements of leaf gas exchange over 2 years showed a significant photosynthetic advantage for the C(4) subspecies under irrigated conditions from spring through autumn. However, the C(4) leaves were killed by winter frost, while photosynthesis continued in the C(3) plants. Unexpectedly, the C(4) 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 C(3) and C(4) grasses.

Consequences of C4 photosynthesis for the partitioning of growth: a test using C3 and C4 subspecies of Alloteropsis semialata under nitrogen-limitation.

C(4) plants dominate the world's subtropical grasslands, but investigations of their ecology typically focus on climatic variation, ignoring correlated changes in soil nutrient concentration. The hypothesis that higher photosynthetic nitrogen use efficiency (PNUE) in C(4) than in C(3) species allows greater flexibility in the partitioning of growth, especially under nutrient-deficient conditions, is tested here. Our experiment applied three levels of N supply to the subtropical grass Alloteropsis semialata, a unique model system with C(3) and C(4) subspecies. Photosynthesis was significantly higher for the same investment of leaf N in the C(4) than C(3) subspecies, and was unaffected by N treatments. The C(4) plants produced more biomass than the C(3) plants at high N levels, diverting a greater fraction of growth into inflorescences and corms, but less into roots and leaves. However, N-limitation of biomass production caused size-dependent shifts in the partitioning of growth. Root production was higher in small than large plants, and associated with decreasing leaf biomass in the C(3), and inflorescence production in the C(4) plants. Higher PNUE in the C(4) than C(3) subspecies was therefore linked with greater investment in sexual reproduction and storage, and the avoidance of N-limitations on leaf growth, suggesting advantages of the C(4) pathway in disturbed and infertile ecosystems.

Drought constraints on C4 photosynthesis: stomatal and metabolic limitations in C3 and C4 subspecies of Alloteropsis semialata.

The C4 photosynthetic pathway uses water more efficiently than the C3 type, yet biogeographical analyses show a decline in C4 species relative to C3 species with decreasing rainfall. To investigate this paradox, the hypothesis that the C4 advantage over C3 photosynthesis is diminished by drought was tested, and the underlying stomatal and metabolic mechanisms of this response determined. The effects of drought and high evaporative demand on leaf gas exchange and photosynthetic electron sinks in C3 and C4 subspecies of the grass Alloteropsis semialata were examined. Plant responses to climatic variation and soil drought were investigated using a common garden experiment with well-watered and natural rainfall treatments, and underlying mechanisms analysed using controlled drying pot experiments. Photosynthetic rates were significantly higher in the C4 than the C3 subspecies in the garden experiment under well-watered conditions, but this advantage was completely lost during a rainless period when unwatered plants experienced severe drought. Controlled drying experiments showed that this loss was caused by a greater increase in metabolic, rather than stomatal, limitations in C4 than in the C3 leaves. Decreases in CO2 assimilation resulted in lower electron transport rates and decreased photochemical efficiency under drought conditions, rather than increased electron transport to alternative sinks. These findings suggest that the high metabolic sensitivity of photosynthesis to severe drought seen previously in several C4 grass species may be an inherent characteristic of the C4 pathway. The mechanism may explain the paradox of why C4 species decline in arid environments despite high water-use efficiency.

Do low standing biomass and leaf area index of sub-tropical coastal dunes ensure that plants have an adequate supply of water?

Water status in relation to standing biomass and leaf area indices (LAI) of the subtropical foredune species Arctotheca populifolia, Ipomoea pes-caprae and Scaevola plumieri were studied in the Eastern Cape, South Africa. The plants showed little evidence of water stress, never developing leaf water potentials more negative than -1.55 MPa, a value which is typical of mesophytes rather than xerophytes. The plants showed no seasonal changes in osmotic potential, an indication that they did not need to osmoregulate, nor were there significant alterations in tissue elasticity. Turgor potential for the most part remained positive throughout the day or recovered positive values at night, a condition suitable for the maintenance of growth that may be essential to cope with sand accretion. All three species show relatively high transpiration rates and only I. pes-caprae showed any evidence of strong limitations of transpiration rate through reductions in midday stomatal conductance. All three species had relatively high instantaneous water use efficiencies as a result of high assimilation rates rather than low transpiration rates. Simple water budgets, accounting for losses by transpiration and inputs from rainfall, suggest that the water stored in the dune sands is sufficient to meet the requirements of the plants, although water budgets calculated for I. pes-caprae suggest that this species may on occasion be water limited. The results suggest that it is the low biomass and LAI that lead to these favourable water relations.