So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

1. Plant Nutrients. 2. The Soil as a Plant Nutrient Medium. 3. Nutrient Uptake and Assimilation. 4. Plant Water Relationships. 5. Plant Growth and Crop Production. 6. Fertilizer Application. 7. Nitrogen. 8. Sulphur. 9. Phosphorus. 10. Potassium. 11. Calcium. 12. Magnesium. 13. Iron. 14. Manganese. 15. Zinc. 16. Copper. 17. Molybdenum. 18. Boron. 19. Further Elements of Importance. 20. Elements with More Toxic Effects. General Readings. References. Index.

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Principles of plant nutrition

At present and more so in the future, irrigated agriculture will take place under water scarcity. Insufficient water supply for irrigation will be the norm rather than the exception, and irrigation management will shift from emphasizing production per unit area towards maximizing the production per unit of water consumed, the water productivity. To cope with scarce supplies, deficit irrigation, defined as the application of water below full crop-water requirements (evapotranspiration), is an important tool to achieve the goal of reducing irrigation water use. While deficit irrigation is widely practised over millions of hectares for a number of reasons—from inadequate network design to excessive irrigation expansion relative to catchment supplies—it has not received sufficient attention in research. Its use in reducing water consumption for biomass production, and for irrigation of annual and perennial crops is reviewed here. There is potential for improving water productivity in many field crops and there is sufficient information for defining the best deficit irrigation strategy for many situations. One conclusion is that the level of irrigation supply under deficit irrigation should be relatively high in most cases, one that permits achieving 60–100% of full evapotranspiration. Several cases on the successful use of regulated deficit irrigation (RDI) in fruit trees and vines are reviewed, showing that RDI not only increases water productivity, but also farmers’ profits. Research linking the physiological basis of these responses to the design of RDI strategies is likely to have a significant impact in increasing its adoption in water-limited areas.

Deficit irrigation for reducing agricultural water use

Osmoregulation and water stress in higher plants

Recent progress in improving the salt tolerance of cultivated plants has been slow. Physiologists have been unable to define single genes or even specific metabolic processes that molecular biologists could target, or pinpoint the part of the plant in which such genes for salt tolerance might be expressed. While the physiological might be expressed. While the physiological processes are undoubtedly complex, faster progress on unraveling mechanisms of salt tolerance might be made if there were more effort to test hypotheses rather than to accumulate data, and to integrate cellular and whole plant responses. This article argues that salts taken up by the plant do not directly control plant growth by affecting turgor, photosynthesis or the activity of any one enzyme. Rather, the build-up of salt in old leaves hasten their death, and the loss of these leaves affects the supply of assimilates or hormones to the growing regions and thereby affects growth.

Physiological processes limiting plant growth in saline soils: some dogmas and hypotheses

Many plant processes are affected by mild water stress, with cell growth probably the most sensitive. Except for turgor-mediated processes, the physicochemical basis for the transduction of small changes in water status into alterations in metabolism remains obscure. Turgor pressure is assigned a critical role in cell growth: the physical force needed to sustain enlargement. Simple physical models relating growth to turgor are conceptually useful in examining effects of water stress but can be misleading because metabolic and regulatory responses may be marked and vary temporally. Osmotic adjustment has long been known as a means by which higher plants adapt to salinity, with much of the cell osmotica being ionic and accumulated from the medium. Though not generally recognized, osmotic adjustment also appears to be an important mechanism for adaptation to water-limiting conditions, even in mesophytic plants. In this case much of the osmotica might possibly be internally generated. Recent field data on seasonal and diurnal adjustment and vertical water-potential gradients in plant canopies are discussed relative to growth and water-potential components.

Water stress, growth, and osmotic adjustment

Plant responses to water deficits, water-use efficiency, and drought resistance

Elongation of intact young leaves of maize was found to be dynamically dependent on soil water supply. With adequate water, elongation was remarkably constant but slowed when the water potential of the soil in pots dropped from -0.1 to -0.2 bar and stopped when it dropped to -2.5 bars. The corresponding range of leaf water potential was -2.8 to -7 bars. Elongation resumed in less than a few seconds after a mildly water-stressed plant was rewatered.The effects on leaf elongation of step-wise changes in water potential of the root solution were determined. When the water potential of the root medium suddenly decreased below 0 bar, growth stopped initially and then resumed at a lower rate. When the water potential was suddenly increased back to 0 bar, growth accelerated transitorily to a high rate before slowing to the steady state rate. These results suggest an increase in cell extensibility during water stress.Leaves stressed for 1 or more days attained after rewatering almost the length of the control leaves. Growth rate after rewatering did not exceed that of the control at the corresponding developmental stage except during the short transitory rapid phase lasting only a fraction of an hour.As stress developed, growth stopped before carbon dioxide assimilation decreased noticeably. Upon the release of mild and short stress, the transitory rapid growth completely made up for the reduced elongation during stress, suggesting that metabolic processes for cell expansion might have proceeded unchecked during the stress period.The sensitivity and rapidity of response to changes in water status all point to the direct role of water in growth; its uptake provides the physical force for cell enlargement.

Immediate and subsequent growth responses of maize leaves to changes in water status.

No-tillage systems affect soil properties depending on the soil, climate, and the time since its implementation. In heavy no-tilled soils a surface compacted layer is commonly found. Such layer can affect root growth and soil water infiltration. In several cases, surface organic carbon can buffer these problems. The aim of this study was to evaluate the effect of 4- and 7-year-old conventional (CT) and no-tillage (NT) treatments on soil physical properties, root growth, and wheat (Triticum turgidum L. var. durum) yield in an Entic Haploxeroll of Central Chile. In both tillage treatments we study soil water retention, bulk density (rb), soil particle density (rs), soil water infiltration, mean-weight diameter of soil aggregates (MWD), penetration resistance, grain yield, and root length density (Lv) up to a depth of 15 cm. The MWD and the penetration resistance were higher under NTas compared to CT. For the top 5 cm of soil, Lv was greater under NTas compared to CT. Differences of Lv between NTand CTwere 2.09, 7.60, and 4.31 cm root cm � 3 soil during the two leaves, flowering and grain filling phenological stages, respectively. Generally, the effect of NTon these properties was more evident near the soil surface. In contrast, fast drainage macropores, rs, and soil water infiltration rates were higher under CT than under NT. Tillage treatments did not significantly affect rb and yield. A longer time under no-tillage enhanced aggregate stability, however, other soil physical properties were negatively affected. # 2008 Elsevier B.V. All rights reserved.

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Soil physical properties and wheat root growth as affected by no-tillage and conventional tillage systems in a Mediterranean environment of Chile

Wild wheat (Triticum aestivum L.) relatives could represent a valuable source of genetic variation for improvement of abiotic stress tolerance in cultivated wheat. A better knowledge of the adaptive strategies developed by these species is needed. A collection of 157 Aegilops geniculata accessions originating from different ecogeographical regions was studied during two successive years for several traits related to water status, chlorophyll content, and plant thermal regulation under Mediterranean field conditions. Close association was found between the studied traits and the origin of accessions. Two adaptive strategies were distinguished. Accessions originating from harsh environments had low biomass, low grain production and high water-use efficiency (low C isotope discrimination). They were early, with small, thick leaves exhibiting low chlorophyll content, high surface temperature and low epidermal transpiration. We suggest that in these accessions, decreased leaf chlorophyll content could limit the energy load from strong sunlight. In accessions originating from regions with a mild Mediterranean climate, thermal regulation of the leaf may rather depend on transpiration, as suggested by high C isotope discrimination values. These accessions also were characterized by high chlorophyll content, leaf area, and biomass production. Associations between the physiological traits observed could help to better understand the relationship between abiotic stress tolerance and yield in cultivated wheats. Results obtained confirmed the potential value of A egilops geniculata for improvement of high temperature and drought stress tolerance in wheat and could contribute to the choice of traits to be introgressed and the accessions to be used in wide hybridization programs.

Edmundo Acevedo

Papers

Water stress, growth, and osmotic adjustment

Plant responses to water deficits, water-use efficiency, and drought resistance

Immediate and subsequent growth responses of maize leaves to changes in water status.

Soil physical properties and wheat root growth as affected by no-tillage and conventional tillage systems in a Mediterranean environment of Chile

Drought and Heat Responses in the Wild Wheat Relative Aegilops geniculata Roth: Potential Interest for Wheat Improvement