Photosynthetic pathway and ecological adaptation explain stomatal trait diversity amongst grasses.

• The evolution of C(4) photosynthesis in plants has allowed the maintenance of high CO(2) assimilation rates despite lower stomatal conductances. This underpins the greater water-use efficiency in C(4) species and their tendency to occupy drier, more seasonal environments than their C(3) relatives. • The basis of interspecific variation in maximum stomatal conductance to water (g(max) ), as defined by stomatal density and size, was investigated in a common-environment screening experiment. Stomatal traits were measured in 28 species from seven grass lineages, and comparative methods were used to test for predicted effects of C(3) and C(4) photosynthesis, annual precipitation and habitat wetness on g(max) . • Novel results were as follows: significant phylogenetic patterns exist in g(max) and its determinants, stomatal size and stomatal density; C(4) species consistently have lower g(max) than their C(3) relatives, associated with a shift towards smaller stomata at a given density. A direct relationship between g(max) and precipitation was not supported. However, we confirmed associations between C(4) photosynthesis and lower precipitation, and showed steeper stomatal size-density relationships and higher g(max) in wetter habitats. • The observed relationships between stomatal patterning, photosynthetic pathway and habitat provide a clear example of the interplay between anatomical traits, physiological innovation and ecological adaptation in plants.

Drought limitation of photosynthesis differs between C3and C4grass species in a comparative experiment.

Phylogenetic analyses show that C4 grasses typically occupy drier habitats than their C3 relatives, but recent experiments comparing the physiology of closely related C3 and C4 species have shown that advantages of C4 photosynthesis can be lost under drought. We tested the generality of these paradoxical findings in grass species representing the known evolutionary diversity of C4 NADP-me and C3 photosynthetic types. Our experiment investigated the effects of drought on leaf photosynthesis, water potential, nitrogen, chlorophyll content and mortality. C4 grasses in control treatments were characterized by higher CO2 assimilation rates and water potential, but lower stomatal conductance and nitrogen content. Under drought, stomatal conductance declined more dramatically in C3 than C4 species, and photosynthetic water-use and nitrogen-use efficiency advantages held by C4 species under control conditions were each diminished by 40%. Leaf mortality was slightly higher in C4 than C3 grasses, but leaf condition under drought otherwise showed no dependence on photosynthetic-type. This phylogenetically controlled experiment suggested that a drought-induced reduction in the photosynthetic performance advantages of C4 NADP-me relative to C3 grasses is a general phenomenon.

Constraining ecosystem processes from tower fluxes and atmospheric profiles.

The planetary boundary layer (PBL) provides an important link between the scales and processes resolved by global atmospheric sampling/modeling and site-based flux measurements. The PBL is in direct contact with the land surface, both driving and responding to ecosystem processes. Measurements within the PBL (e.g., by radiosondes, aircraft profiles, and flask measurements) have a footprint, and thus an integrating scale, on the order of 1-100 km. We use the coupled atmosphere-biosphere model (CAB) and a Bayesian data assimilation framework to investigate the amount of biosphere process information that can be inferred from PBL measurements. We investigate the information content of PBL measurements in a two-stage study. First, we demonstrate consistency between the coupled model (CAB) and measurements, by comparing the model to eddy covariance flux tower measurements (i.e., water and carbon fluxes) and also PBL scalar profile measurements (i.e., water, carbon dioxide, and temperature) from Canadian boreal forest. Second, we use the CAB model in a set of Bayesian inversions experiments using synthetic data for a single day. In the synthetic experiment, leaf area and respiration were relatively well constrained, whereas surface albedo and plant hydraulic conductance were only moderately constrained. Finally, the abilities of the PBL profiles and the eddy covariance data to constrain the parameters were largely similar and only slightly lower than the combination of both observations.

Responses of global plant diversity capacity to changes in carbon dioxide concentration and climate.

We model plant species diversity globally by country to show that future plant diversity capacity has a strong dependence on changing climate and carbon dioxide concentration. CO2 increase, through its impact on net primary production and warming is predicted to increase regional diversity capacity, while warming with constant CO2 leads to decreases in diversity capacity. Increased CO2 concentrations are unlikely to counter projected extinctions of endemic species, shown in earlier studies to be more strongly dependent on changing land use patterns than climate per se. Model predictions were tested against (1) contemporary observations of tree species diversity in different biomes, (2) an independent global map of contemporary species diversity and (3) time sequences of plant naturalisation for different locations. Good agreements between model, observations and naturalisation patterns support the suggestion that future diversity capacity increases are likely to be filled from a 'cosmopolitan weed pool' for which migration appears to be an insignificant barrier.

Constraining the Sheffield dynamic global vegetation model using stream-flow measurements in the United Kingdom.

The biospheric water and carbon cycles are intimately coupled, so simulating carbon fluxes by vegetation also requires modelling of the water fluxes, with each component influencing the other. Observations of river streamflow integrate information at the catchment scale and are widely available over a long period; they therefore provide an important source of information for validating or calibrating vegetation models. In this paper, we analyse the performance of the Sheffield dynamic global vegetation model (SDGVM) for predicting river streamflow and quantifying how this information helps to constrain carbon flux predictions. The SDGVM is run for 29 large catchments in the United Kingdom. Annual streamflow estimates are compared with long time-series observations. In 23 out of the 29 catchments, the bias between model and observations is less than 50 mm, equivalent to less than 10% of precipitation. In the remaining catchments, larger errors are because of combinations of unpredictable causes, in particular various human activities and measurement issues and, in two cases, unidentified causes. In one of the catchments, we assess to what extent a knowledge of annual streamflow can constrain model parameters and in turn constrain estimates of gross primary production (GPP). For this purpose, we assume the model parameters are uncertain and constrain them by the streamflow observations using the generalized likelihood uncertainty estimation method. Comparing the probability density function of GPP with and without constraint shows that streamflow effectively constrains GPP, mainly by setting a low probability to GPP values below about 1100 g C-1 m2 yr-1 . In other words, streamflow observations allow the rejection of low values of GPP, so that the potential range of possible GPP values is almost halved.

Predicting plant responses to global environmental change.

Predicting the future responses of plants and ecosystems to further changes in the CO2 concentration of the atmosphere and to the possibility of global warming are important current concerns. Predictions have been most frequently attempted using short-term, single-factor experiments in controlled environments. However, these experiments have failed to indicate the outcome of field experiments at larger spatial and temporal scales. Some of this failure is due to ignorance of environmental conditions and interactions while some is due to the use of inappropriate short-cuts, such as the addition of fertilizers for simulating enhanced mineralization, and some is due to ignorance of the processes involved in scaling-up from individual plants to populations. Long-term observations on plants in ecosystems nevertheless indicate that community-scale experiments may provide a useful but imperfect capacity to predict ecosystem responses. Although difficult to implement in practice, it is concluded that catchment-scale experiments offer the best opportunity to predict plant, community and ecosystem responses to environmental change. CONTENTS Summary 239 I. Introduction 239 II. CO2 -enrichment experiments 240 III. Experiments with applied nitrogen fertilizer 243 IV. Population ecophysiology 245 V. Ecosystem predictions 248 Acknowledgements 249 References 250.

Ecophysiological responses of plants to global environmental change since the Last Glacial Maximum.

Ecophysiological information on the responses of plants to past global environmental changes may be obtained from Quaternary fossil leaves by measurements of (i) stomatal density, (ii) stomatal dimensions and (iii) 13 C discrimination (Δ13 C). The stomatal density and stomatal dimensions of leaves can be used to calculate stomatal conductance, while leaf Δ13 C values provide independent information on stomatal conductance and plant water use efficiency. In this paper, stomatal conductance is calculated for a sequence of radiocarbon dated fossil leaves of Salix herbacea L. which, together with herbarium and fresh material, represents a time-series spanning from the Last Glacial Maximum (LGM) (16500 yr BP) to the present day. The calculated values were then tested against leaf Δ13 C values previously reported for the same material. Our calculations show that stomatal conductance is negatively correlated with increases in atmospheric CO2 concentration over the last 16500 yr. This represents the first evidence of long-term response of stomatal conductance to increases in atmospheric CO2 concentration and confirms the response observed in experimental systems exposing plants to lower-than-present CO2 concentrations in controlled environments. The calculated decrease in conductance was positively correlated with leaf Δ13 C values, supporting this interpretation. The mean leaf Δ13 C value for the 18th and 19th centuries was significantly (P≥ 0.05) lower than the mean for the interval LGM-Holocene (10000 yr BP) implying an increase in plant water-use-efficiency over this time. These two lines of evidence, together with the stomatal density record from a glacial cycle, and experimental studies growing C3 plants in glacial-to-present CO2 concentrations, strongly imply that the water use efficiency of vegetation during the LGM was lower than at present and that it has increased since that time. Further evidence in support of this conclusion comes from the pattern of world vegetation types present during the LGM previously reconstructed using palaeoecological data. This evidence demonstrates that the distribution of vegetation types during the LGM was significantly different from that of the present day and showed a contraction in the area of rain forest and a major expansion of desert areas.

Stomatal development and CO2 : ecological consequences.

• Stomatal density responses by 48 accessions of Arabidopsis, to CO2 enrichment, broadly parallel interspecific observations. • Accessions differing in the degree of stomatal response to both CO2 and drought differed in flower production. Under well watered conditions flowering benefits from a small reduction in stomatal density with CO2 enrichment, but benefits from a large reduction under drought. • Stomatal density increases with altitude in Vaccinium myrtillus but is also strongly influenced by exposure. Exposed plants had higher stomatal densities than plants at the same altitude but in a community of individuals. This difference might be explained by systemic signalling within the plant as mature leaves detect both irradiance and [CO2 ], subsequently controlling the response of stomatal development in developing leaves. • Plants with the highest stomatal densities also had the highest stomatal conductances and photosynthetic rates. This suggests that signalling from mature to developing leaves predetermines the potential of the developing leaf to maximize its photosynthetic potential, including associated features such as nitrogen allocation, during early stages of development in the enclosed bud.

The Lowland-to-Upland Transition--Modelling Plant Responses to Environmental Change.

A published correlative model has predicted that the distributional limits of plants and vegetation zones on mountains will increase in altitude with global warming. I test this hypothesis using results from published experimental studies. Investigations and models of the responses of leaf growth to temperature are in accord with the prediction. However, the individualistic responses of species to CO2 enrichment indicate that the prediction is unlikely to be true for all species: growth is stimulated by CO2 enrichment for some species but not for others. Wind speed generally increases with altitude on mountains, and plants from high altitude tend to be more wind resistant than species from the lowland. Therefore it is expected that, particularly on wind-swept mountains, global warming will not necessarily be followed by the spread of lowland species into the uplands.

Ecophysiological studies on the shrub Vaccinium myrtillus L. taken from a wide altitudinal range.

Observations have been made on the gas exchange and morphology of Vaccinium myrtillus taken from altitudes of 200 m, 610 m and 1,100 m along an altitudinal gradient in central Scotland. Under saturating irradiance, optimum temperatures and a range of vapour pressure deficits, photosynthetic rate and stomatal conductance increased with the altitude of origin of the populations. Correlated with these increases was an increase in the adaxial stomatal density with altitude. This response to altitude could be simulated in controlled conditions, by growing plants in a CO2 concentration below ambient, similar to that expected at altitude.Plant height decreased with altitude, a feature which was maintained in cultivation. Stem rigidity declined with altitude, in a manner which is predicted to limit the reproductive capacity of the population from 1,100 m in high wind speeds.Total leaf nitrogen increased with altitude. The nitrogen economy of the shoot is discussed in terms of nitrogen availability for stems and leaves and its control over maximum rates of photosynthesis, competitive ability and reproductive capacity.

The influences of increased CO2 and water supply on growth, biomass allocation and water use efficiency of Sinapis alba L. grown under different wind speeds.

We examined how independent and interactive effects of CO2 concentrations, water supply and wind speed affect growth rates, biomass partitioning, water use efficiency, diffusive conductance and stomatal density of plants. To test the prediction that wind stress will be ameliorated by increased CO2 and/or by unrestricted water supply we grew Sinapis alba L. plants in controlled chambers under combinations of two levels of CO2 (350 ppmv, 700 ppmv), two water regimes and two wind speeds (0.3 ms-1, 3.7 ms-1). We harvested at ten different dates over a period of 60 days. A growth analysis was carried out to evaluate treatment effects on plant responses. Plants grown both in increased CO2 and in low wind conditions had significantly greater stem length, leaf area and dry weights of plant parts. Water supply significantly affected stem diameter, root weight and leaf area. CO2 enrichment significantly increased the rate of biomass accumulation and the relative ratio of biomass increase to leaf area expansion. High wind speed significantly reduced plant growth rates and the rate of leaf area expansion was reduced more than the rate of biomass accumulation. Regression analysis showed significant CO2 effects on the proportion of leaf and stem dry weight to total dry weight. A marked plant-age effect was dependent on water supply, wind speed and CO2 concentration. A reduced water supply significantly decreased the stomatal conductance, and water use efficiency significantly increased with a limited water supply, low wind and increased CO2. We found significant CO2 x wind effects for water diffusion resistance, adaxial number of stomata and water use efficiencies and significant wind x water effect for water use efficiency. In conclusion, wind stress was ameliorated by growing in unrestricted water but not by growing in increased CO2.

The dynamics of leaf extension in plants with diverse altitudinal ranges : II. Field studies in Poa species between 600 and 3200 m altitude.

The dynamics of leaf extension in five species of Poa were studied with electronic auxanometers (LVDTs) along an elevational gradient from 600 to 3200 m in the Austrian Alps. Extension rates peak at midday at all elevations and rates at 20°C are almost twice as high at low elevation as compared with those from the highest sites. The low temperature threshold for leaf extension drops by 7 K over this range of elevation, with plants from the highest sites showing some extension around freezing point. Thus, there is a substantial adaptive adjustment in response of leaf extension to declining mean temperatures with increasing altitude, which is not paralleled by known altitudinal trends of photosynthetic responses in herbaceous plants.

The influence of plant density on the responses of Sinapis alba to CO2 and windspeed.

Plants in nature live in populations of variable density, a characteristic which may influence individual plant responses to the environment. We investigated how the responses of Sinapis alba plants to different wind speeds and CO2 concentrations could be modified by plant density. In our wind-density experiment the expectation that mechanical and physiological effects of wind will be ameliorated by growing in high density, as a result of positive plant interactions, was realised. Although individual plants were smaller at higher densities, the effect of increasing windspeed was much less than at lower plant densities. A similar reduced sensitivity of individual plant growth under high densities was also observed under CO2 enrichment. When measured as a population or stand response, there was no effect of density on the CO2 responses, with all stands showing very similar increases in total biomass with CO2 enrichment. In the wind speed experiment, total biomass per stand increased significantly with density, although there was no effect of density on the wind speed response. Specific leaf area decreased with increasing wind speed and this response was significantly affected by the density at which the plants grew.

The dynamics of leaf extension in plants with diverse altitudinal ranges : I. Field observations on temperature responses at one altitude.

Rates of leaf extension have been studied with electronic auxanometers at mid-altitude in the Austrian Alps, where both low and high altitude species co-occur. The results demonstrate a clear differentiation in the temperature responses of extension between these two groups of species. For the low or mid-altitude species of Achillea millefolium, Agrostis stolonifera, Poa alpina and Rumex arifolius, the average rate of leaf extension increases from 0.1 to 0.4 mm h-1 between 10 and 20° C. For the high-alpine species of Achillea erba-rotta ssp moschata, Poa alpina ssp vivipara and Polygonum viviparum the average rate of leaf extension was considerably lower from 0.016 to 0.064 mm h-1, between 10 and 20° C.Leaf extension in the lowland species was not observed below an average temperature of about 5° C, whilst no limit was observed for the upland species, down to a temperature of about 0° C.In the cases of the dicotyledons that were studied, leaf plus petiole shrinkage was observed to occur, for as much as 2 to 4 h, during periods of high water vapour pressure deficits. This response was not observed for the monocotyledons.The observations of leaf extension show that daily totals of extension in species from high altitudies will be much less sensitive to day, to day variations in local climate than will the species from low altitudes. The lowland species will have higher rates of extension during clear and warm weather conditions but lower rates in cold, cloudy weather.