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Biomedical subjects

F I Woodward

Publications and source records attributed to F I Woodward.

10 recordsLinked to original sources

Systemic signalling of environmental cues in Arabidopsis leaves.

Light intensity and atmospheric CO2 partial pressure are two environmental signals known to regulate stomatal numbers. It has previously been shown that if a mature Arabidopsis leaf is supplied with either elevated CO2 (750 ppm instead of ambient at 370 ppm) or reduced light levels (50 micromol m-2 s-1 instead of 250 micromol m-2 s-1), the young, developing leaves that are not receiving the treatment grow with a stomatal density as if they were exposed to the treatment. But the signal(s) that it is believed is generated in the mature leaves and transmitted to developing leaves are largely unknown. Photosynthetic rates of treated, mature Arabidopsis leaves increased in elevated CO2 and decreased when shaded, as would be expected. Similarly, the levels of sugars (glucose, fructose, and sucrose) in the treated mature leaves increased in elevated CO2 and decreased with shade treatment. The levels of sugar in developing leaves were also measured and it was found that they mirrored this result even though they were not receiving the shade or elevated CO2 treatment. To investigate the effect of these treatments on global gene expression patterns, transcriptomics analysis was carried out using Affymetrix, 22K, and ATH1 arrays. Total RNA was extracted from the developing leaves after the mature leaves had received either the ambient control treatment, the elevated CO2 treatment, or the shade treatment, or both elevated CO2 and shade treatments for 2, 4, 12, 24, 48, or 96 h. The experiment was replicated four times. Two other experiments were also conducted, one to compare and contrast gene expression in response to plants grown at elevated CO2 and the other to look at the effect of these treatments on the mature leaf. The data were analysed and 915 genes from the untreated, signalled leaves were identified as having expression levels affected by the shade treatment. These genes were then compared with those whose transcript abundance was affected by the shade treatment in the mature treated leaves (1181 genes) and with 220 putative 'stomatal signalling' genes previously identified from studies of the yoda mutant. The results of these experiments and how they relate to environmental signalling are discussed, as well as possible mechanisms for systemic signalling.

Acclimatization↗

The global distribution of ecosystems in a world without fire.

This paper is the first global study of the extent to which fire determines global vegetation patterns by preventing ecosystems from achieving the potential height, biomass and dominant functional types expected under the ambient climate (climate potential). To determine climate potential, we simulated vegetation without fire using a dynamic global-vegetation model. Model results were tested against fire exclusion studies from different parts of the world. Simulated dominant growth forms and tree cover were compared with satellite-derived land- and tree-cover maps. Simulations were generally consistent with results of fire exclusion studies in southern Africa and elsewhere. Comparison of global 'fire off' simulations with landcover and treecover maps show that vast areas of humid C(4) grasslands and savannas, especially in South America and Africa, have the climate potential to form forests. These are the most frequently burnt ecosystems in the world. Without fire, closed forests would double from 27% to 56% of vegetated grid cells, mostly at the expense of C(4) plants but also of C(3) shrubs and grasses in cooler climates. C(4) grasses began spreading 6-8 Ma, long before human influence on fire regimes. Our results suggest that fire was a major factor in their spread into forested regions, splitting biotas into fire tolerant and intolerant taxa.

Biological Evolution↗

Global climate and the distribution of plant biomes.

Biomes are areas of vegetation that are characterized by the same life-form. Traditional definitions of biomes have also included either geographical or climatic descriptors. This approach describes a wide range of biomes that can be correlated with characteristic climatic conditions, or climatic envelopes. The application of remote sensing technology to the frequent observation of biomes has led to a move away from the often subjective definition of biomes to one that is objective. Carefully characterized observations of life-form, by satellite, have been used to reconsider biome classification and their climatic envelopes. Five major tree biomes can be recognized by satellites based on leaf longevity and morphology: needleleaf evergreen, broadleaf evergreen, needleleaf deciduous, broadleaf cold deciduous and broadleaf drought deciduous. Observations indicate that broadleaf drought deciduous vegetation grades substantially into broadleaf evergreen vegetation. The needleleaf deciduous biome occurs in the world's coldest climates, where summer drought and therefore a drought deciduous biome are absent. Traditional biome definitions are quite static, implying no change in their life-form composition with time, within their particular climatic envelopes. However, this is not the case where there has been global ingress of grasslands and croplands into forested vegetation. The global spread of grasses, a new super-biome, was probably initiated 30-45 Myr ago by an increase in global aridity, and was driven by the natural spread of the disturbances of fire and animal grazing. These disturbances have been further extended over the Holocene era by human activities that have increased the land areas available for domestic animal grazing and for growing crops. The current situation is that grasses now occur in most, if not all biomes, and in many areas they dominate and define the biome. Croplands are also increasing, defining a new and relatively recent component to the grassland super-biome. In the case of both grassland and croplands, various forms of disturbance, particularly frequent disturbance, lead to continued range extensions of the biomes.

Climate↗

Vegetation dynamics--simulating responses to climatic change.

A modelling approach to simulating vegetation dynamics is described, incorporating critical processes of carbon sequestration, growth, mortality and distribution. The model has been developed to investigate the responses of vegetation to environmental change, at time scales from days to centuries and from the local to the global scale. The model is outlined and subsequent tests, against independent data sources, are relatively successful, from the small scale to the global scale. Tests against eddy covariance observations of carbon exchange by vegetation indicated significant differences between measured and simulated net ecosystem production (NEP). NEP is the net of large fluxes due to gross primary production and respiration, which are not directly measured and so there is some uncertainty in explaining differences between observations and simulations. In addition it was noted that closer agreement of fluxes was achieved for natural, or long-lived managed vegetation than for recently managed vegetation. The discrepancies appear to be most closely related to respiratory carbon losses from the soil, but this area needs further exploration. The differences do not scale up to the global scale, where simulated and measured global net biome production were similar, indicating that fluxes measured at the managed observed sites are not typical globally. The model (the Sheffield Dynamic Global Vegetation Model, SDGVM) has been applied to contemporary vegetation dynamics and indicates a significant CO2 fertilisation effect on the sequestration of atmospheric CO2. The terrestrial carbon sink for the 20th century is simulated to be widespread between latitudes 40 degrees S and 65 degrees N, but is greatest between 10 degrees S and 6 degrees N, excluding the effects of human deforestation. The mean maximum sink capacity over the 20th century is small, at 25 gC m(-2) year(-1), or approximately 1% of gross primary production. Simulations of vegetation dynamics under a scenario of future global warming indicate a gradual decline in the terrestrial carbon sink, with the capacity to absorb human emissions of CO2 being reduced from 20% in 2000 to approximately 2% between 2075 and 2100. The responses of carbon sequestration and vegetation structure and distribution to stabilisation of climate and CO2 may extend for up to 50 years after stabilisation has occurred.

Adaptation, Physiological↗

The HIC signalling pathway links CO2 perception to stomatal development.

Stomatal pores on the leaf surface control both the uptake of CO2 for photosynthesis and the loss of water during transpiration. Since the industrial revolution, decreases in stomatal numbers in parallel with increases in atmospheric CO2 concentration have provided evidence of plant responses to changes in CO2 levels caused by human activity. This inverse correlation between stomatal density and CO2 concentration also holds for fossil material from the past 400 million years and has provided clues to the causes of global extinction events. Here we report the identification of the Arabidopsis gene HIC (for high carbon dioxide), which encodes a negative regulator of stomatal development that responds to CO2 concentration. This gene encodes a putative 3-keto acyl coenzyme A synthase--an enzyme involved in the synthesis of very-long-chain fatty acids. Mutant hic plants exhibit up to a 42% increase in stomatal density in response to a doubling of CO2. Our results identify a gene involved in the signal transduction pathway responsible for controlling stomatal numbers at elevated CO2.

Algal Proteins↗

Plant diversity and productivity experiments in european grasslands

At eight European field sites, the impact of loss of plant diversity on primary productivity was simulated by synthesizing grassland communities with different numbers of plant species. Results differed in detail at each location, but there was an overall log-linear reduction of average aboveground biomass with loss of species. For a given number of species, communities with fewer functional groups were less productive. These diversity effects occurred along with differences associated with species composition and geographic location. Niche complementarity and positive species interactions appear to play a role in generating diversity-productivity relationships within sites in addition to sampling from the species pool.

Journal Article↗

Temperature and the distribution of plant species.

An understanding of the mechanisms by which temperature influences the distribution of species and vegetation has been attempted by modelling population growth and establishing those stages of the plant life cycle which, when diminished by extremes of temperature, for example, may have the greatest impact on plant survival. This analysis suggests that the heat sum of the growing season, measured as day-degrees, controls the distribution of annual vegetation. For perennial vegetation both the heat sum of the growing season and the annual, absolute minimum temperature are critical. Climatic correlations and experimental analyses indicate that, in northern Europe, the northern latitudinal and upper altitudinal limits of lowland and southern vegetation are directly controlled by climate. In contrast, the southern and lower altitudinal limits of upland and northern vegetation are likely to be controlled by temperature-sensitive competition with southern or lowland species. Many of the temperature-sensitive processes of plant growth and development, such as the non-linearity of extension growth and variations in the threshold temperatures of processes, may increase the realized heat sum at a particular geographical location. However, in more northerly climates, photoperiodic control is crucial in avoiding precocious development in the highly variable climatic conditions of early spring.

Adaptation, Physiological↗

Phosphoenolpyruvate carboxylase from pennywort (Umbilicus rupestris). Changes in properties after exposure to water stress.

Umbilicus rupestris (pennywort) switches from C3 photosynthesis to an incomplete form of crassulacean acid metabolism (referred to as 'CAM-idling') when exposed to water stress (drought). This switch is accompanied by an increase in the activity of phosphoenolpyruvate carboxylase. This enzyme also shows several changes in properties, including a marked decrease in sensitivity to acid pH, a lower Km for phosphoenolpyruvate, very much decreased sensitivity to the allosteric inhibitor malate, and increased responsiveness to the allosteric effector glucose 6-phosphate. The Mr of the enzyme remains unchanged, at approx. 185 000. These changes in properties of phosphoenolpyruvate carboxylase are discussed in relation to the roles of the enzyme in C3 and in CAM plants.

Carboxy-Lyases↗