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Stomatal functioning regulates the fluxes of CO2 and water vapor between vegetation and atmosphere and thereby influences plant adaptation to their habitats. Stomatal traits are controlled by external environmental and internal cellular signaling. The objective of this study was to quantify the effects of CO2 enrichment (CE) on stomatal density (SD)-related properties, guard cell length (GCL) and carbon isotope ratio (δ13C) of a range of Arabidopsis thaliana ecotypes originating from a wide altitudinal range [50-1260 m above sea level (asl)], and grown at 400 and 800 ppm [ CO2], and thereby elucidate the possible adaptation and acclimation responses controlling stomatal traits and water use efficiency (WUE). There was a highly significant variation among ecotypes in the magnitude and direction of response of stomatal traits namely, SD and stomatal index (SI) and GCL, and δ13C to CE, which represented a short-term acclimation response. A majority of ecotypes showed increased SD and SI with CE with the response not depending on the altitude of origin. Significant ecotypic variation was shown in all stomatal traits and δ13C at each [ CO2]. At 400 ppm, means of SD, SI and GCL for broad altitudinal ranges, i.e. low (<100 m), mid (100-400 m) and high (>400 m), increased with increasing altitude, which represented an adaptation response to decreased availability of CO2 with altitude. δ13C was negatively correlated to SD and SI at 800 ppm but not at 400 ppm. Our results highlight the diversity in the response of key stomatal characters to CE and altitude within the germplasm of A. thaliana and the need to consider this diversity when using A. thaliana as a model plant. [ABSTRACT FROM AUTHOR]

Summary [ABSTRACT FROM AUTHOR]

• The recent discovery of a strong positive relationship between angiosperm genome size and stomatal guard cell length (GCL) opens the possibility of using plant fossil guard cell size as a proxy for changes in angiosperm genome size over periods of environmental change. • The responses of GCL to environmental stimuli are currently unknown and may obscure this predictive relationship. • Here, we investigated the effects of environmental variables (atmospheric CO2, drought, relative humidity, irradiance, ultraviolet radiation and pathogen attack) on GCL in the model plant Arabidopsis thaliana to quantify environmentally induced variation. • GCL responded to all variables tested, but the changes incurred did not significantly impinge on the predictive capability of the relationship. [ABSTRACT FROM AUTHOR]

• The observation that stomatal density (number mm−2) on herbarium leaves had decreased over the last century represents clear evidence that plants have responded to anthropogenic increases in CO2 concentration. The mechanism of the response has proved elusive but here it is shown that density responses to both CO2 concentration and humidity are correlated with changes in whole-plant transpiration and leaf abscisic acid (ABA) concentration. • The transpiration rate of a range of accessions of Arabidopsis thaliana was manipulated by changing CO2 concentration, humidity and by exogenous application of ABA. Stomatal density increased with transpiration and leaf ABA concentration. • A common property of signal transduction systems is that they rapidly lose their ability to respond to the co-associated stimulus. Pathways of water movement within the plant are connected and so variations in supply and demand can be signalled throughout the plant directly, modifying stomatal aperture of mature leaves and stomatal density of developing leaves. Furthermore, the system identified here does not conform to the loss of ability to respond. • A putative mechanism is proposed for the control of stomatal density by transpiration rate and leaf ABA concentration. [ABSTRACT FROM AUTHOR]

Stomata, the small pores on the surfaces of leaves and stalks, regulate the flow of gases in and out of leaves and thus plants as a whole. They adapt to local and global changes on all timescales from minutes to millennia. Recent data from diverse fields are establishing their central importance to plant physiology, evolution and global ecology. Stomatal morphology, distribution and behaviour respond to a spectrum of signals, from intracellular signalling to global climatic change. Such concerted adaptation results from a web of control systems, reminiscent of a ‘scale-free’ network, whose untangling requires integrated approaches beyond those currently used. [ABSTRACT FROM AUTHOR]

Summary • 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. [ABSTRACT FROM AUTHOR]

Stomatal numbers are tightly controlled by environmental signals including light intensity and atmospheric CO2 partial pressure. This requires control of epidermal cell development during the early phase of leaf growth and involves changes in both the density of cells on the leaf surface and the proportion of cells that adopt a stomatal fate. This paper reviews the current understanding of how stomata develop and describes recent advances that have given insights into the regulatory mechanisms involved using mutant Arabidopsis plants that implicates a role for long-chain fatty acids in cell-to-cell communication. Evidence is presented which indicates that long-distance signalling from mature to newly developing leaves forms part of the mechanism by which stomatal development responds to environmental cues. Analysis of mutant plants suggests that the plant hormones abscisic acid, ethylene and jasmonates are implicated in the long-distance signalling pathway and that the action may be mediated by reactive oxygen species. [ABSTRACT FROM AUTHOR]

Reports the identification of the Arabidopsis gene HIC (for high carbon dioxide), which encodes a negative regulator of stomatal development that responds to carbon dioxide concentration. Information of the synthase it produces, which is an enzyme involved in the synthesis of very-long-chain fatty acids; Results of experiments done on mutant hic plants.

A survey of 100 species and 122 observations has shown an average reduction in stomatal density of 143% (SE ± 2.2%) with CO2 enrichment, with 74% of the cases exhibiting a reduction in stomatal density. A sign test demonstrated that stomata! density decreases, in response to CO2, significantly more often than expected by chance. Repeated observations on the same species indicated a significant repeatability in the direction of the stomatal response. Analyses which removed the potential effect of taxonomy on this data set showed no significant patterns in the dependency of the degree of stomatal change on growth form (woodiness vs. non-woodiness; trees vs. shrubs), habitat (cool VS. warm) or stomatal distribution on the leaf (amphi- vs. hypostomatous). Forty-three of the observations had been made in controlled environments and under a typical range in CO2 enrichment of 350-700 μmol mol-1. For these cases the average stomata! density declined by 9% (SE ± 3.3%) and 60% of the cases showed reductions in stomatal density. When analyses were restricted to these 43 observations, amphistomatous samples more frequently had greater changes in stomatal density than did hypostomatous samples. The degree of reduction in stomatal density with increasing CO2 increases with initial stomatal density, after the influence of taxonomy is removed using analyses of independent contrasts. When the data were examined for patterns that might be due explicitly to the effects of relatedness, the subclasses of the Hamamelidae and the Rosidae showed highly significant reductions in stomatal density with CO2 (87% of the specks studied in the Hamamelidae and 80% of the species in the Rosidae showed reduction with CO2 enrichment) and correlations between initial stomatal density and degree of reduction in stomatal density. [ABSTRACT FROM AUTHOR]

Ecophysiological information on the responses of plants to past global environment changes may be obtained from quaternary fossil leaves by measurements of (i) stomatal density, (ii) stomatal dimensions and (iii) 13C discrimination (Δ 13C) The stomata! density and stomatal dimensions of leaves can be used to calculate stomatal conductance, while leaf Δ 13C values provide independent information on stomata! 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 Δ 13C 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 stomata! 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 δ 13C values, supporting this interpretation The mean leaf δ 13C value for the 18th and 19th centuries was significantly (P < 005) 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 horn 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. [ABSTRACT FROM AUTHOR]

It is sometimes assumed that because increases in atmospheric CO2 concentration usually enhance water use efficiency per unit leaf area, there will be a tendency for plants to show greater drought tolerance as well as increased biomass in the future. A critical examination of the responses to elevated CO2 in three temperate tree species shows that this assumption might be incorrect in the case of two of them. Both beech (Fagus sylvatica L.) and birch (Betula pubescens Ehrh.) display minimal stomatal closing responses to elevated CO2, and in the case of F. sylvatica the stomatal control of transpiration per unit leaf area appears to be unable to compensate for the greater development of leaf area. By contrast, the stomata of oak (Quercus robur L.) close appreciably in elevated CO2, to an extent which might be sufficient to compensate for an increase in total leaf area. A simple model for the controls on water supply and consumption for the whole tree suggests that in F. sylvatica the potential height attainment for a given sapwood area might decrease as the atmospheric CO2 concentration rises. The conclusions drawn from experimental data and from modelling are supported by field observations made in the UK in 1995, when the three species responded very differently to severe drought. We suggest that the progressive increase in the concentration of atmospheric CO2 over the past 200 yr might have accentuated differences in drought sensitivity between these species. [ABSTRACT FROM AUTHOR]

The article introduces research on the variety of mechanisms by which plants adjust to variations in water supply, including one on aquaporins and another on the role of stomatal responses in rapid drought responses.