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

Scarcity of water is a severe environmental constraint to plant productivity. Drought-induced loss in crop yield probably exceeds losses from all other causes, since both the severity and duration of the stress are critical. Here, we have reviewed the effects of drought stress on the growth, phenology, water and nutrient relations, photosynthesis, assimilate partitioning, and respiration in plants. This article also describes the mechanism of drought resistance in plants on a morphological, physiological and molecular basis. Various management strategies have been proposed to cope with drought stress. Drought stress reduces leaf size, stem extension and root proliferation, disturbs plant water relations and reduces water-use efficiency. Plants display a variety of physiological and biochemical responses at cellular and whole-organism levels towards prevailing drought stress, thus making it a complex phenomenon. CO2 assimilation by leaves is reduced mainly by stomatal closure, membrane damage and disturbed activity of various enzymes, especially those of CO2 fixation and adenosine triphosphate synthesis. Enhanced metabolite flux through the photorespiratory pathway increases the oxidative load on the tissues as both processes generate reactive oxygen species. Injury caused by reactive oxygen species to biological macromolecules under drought stress is among the major deterrents to growth. Plants display a range of mechanisms to withstand drought stress. The major mechanisms include curtailed water loss by increased diffusive resistance, enhanced water uptake with prolific and deep root systems and its efficient use, and smaller and succulent leaves to reduce the transpirational loss. Among the nutrients, potassium ions help in osmotic adjustment; silicon increases root endodermal silicification and improves the cell water balance. Low-molecular-weight osmolytes, including glycinebetaine, proline and other amino acids, organic acids, and polyols, are crucial to sustain cellular functions under drought. Plant growth substances such as salicylic acid, auxins, gibberrellins, cytokinin and abscisic acid modulate the plant responses towards drought. Polyamines, citrulline and several enzymes act as antioxidants and reduce the adverse effects of water deficit. At molecular levels several drought-responsive genes and transcription factors have been identified, such as the dehydration-responsive element-binding gene, aquaporin, late embryogenesis abundant proteins and dehydrins. Plant drought tolerance can be managed by adopting strategies such as mass screening and breeding, marker-assisted selection and exogenous application of hormones and osmoprotectants to seed or growing plants, as well as engineering for drought resistance.

https://hal.archives-ouvertes.fr/hal-00886451/document

Plant drought stress: effects, mechanisms and management

There is growing recognition that classifying terrestrial plant species on the basis of their function (into 'functional types') rather than their higher taxonomic identity, is a promising way forward for tackling important ecological questions at the scale of ecosystems, landscapes or biomes. These questions include those on vegetation responses to and vegetation effects on, environmental changes (e.g. changes in climate, atmospheric chemistry, land use or other disturbances). There is also growing consensus about a shortlist of plant traits that should underlie such functional plant classifications, because they have strong predictive power of important ecosystem responses to environmental change and/or they themselves have strong impacts on ecosystem processes. The most favoured traits are those that are also relatively easy and inexpensive to measure for large numbers of plant species. Large international research efforts, promoted by the IGBP–GCTE Programme, are underway to screen predominant plant species in various ecosystems and biomes worldwide for such traits. This paper provides an international methodological protocol aimed at standardising this research effort, based on consensus among a broad group of scientists in this field. It features a practical handbook with step-by-step recipes, with relatively brief information about the ecological context, for 28 functional traits recognised as critical for tackling large-scale ecological questions.

/pdf/a-handbook-of-protocols-for-standardised-and-easy-2uo4112zhh.pdf

A handbook of protocols for standardised and easy measurement of plant functional traits worldwide

Heat tolerance in plants: An overview

Plant functional traits are the features (morphological, physiological, phenological) that represent ecological strategies and determine how plants respond to environmental factors, affect other trophic levels and influence ecosystem properties. Variation in plant functional traits, and trait syndromes, has proven useful for tackling many important ecological questions at a range of scales, giving rise to a demand for standardised ways to measure ecologically meaningful plant traits. This line of research has been among the most fruitful avenues for understanding ecological and evolutionary patterns and processes. It also has the potential both to build a predictive set of local, regional and global relationships between plants and environment and to quantify a wide range of natural and human-driven processes, including changes in biodiversity, the impacts of species invasions, alterations in biogeochemical processes and vegetation–atmosphere interactions. The importance of these topics dictates the urgent need for more and better data, and increases the value of standardised protocols for quantifying trait variation of different species, in particular for traits with power to predict plant- and ecosystem-level processes, and for traits that can be measured relatively easily. Updated and expanded from the widely used previous version, this handbook retains the focus on clearly presented, widely applicable, step-by-step recipes, with a minimum of text on theory, and not only includes updated methods for the traits previously covered, but also introduces many new protocols for further traits. This new handbook has a better balance between whole-plant traits, leaf traits, root and stem traits and regenerative traits, and puts particular emphasis on traits important for predicting species’ effects on key ecosystem properties. We hope this new handbook becomes a standard companion in local and global efforts to learn about the responses and impacts of different plant species with respect to environmental changes in the present, past and future.

/pdf/new-handbook-for-standardised-measurement-of-plant-5aljxhhg5y.pdf

New handbook for standardised measurement of plant functional traits worldwide

This presentation is a concept review paper dealing with a central dilemma in understanding, designing, and acting upon crop plant improvement programs for drought conditions. The association among yield potential (YP), drought resistance (DR), and water-use efficiency (WUE) is often misunderstood, which in turn can lead to conceptual oversight and wrong decisions in implementing breeding programs for drought-prone environments. Although high YP is the target of most crop breeding programs, it might not be compatible with superior DR. On the other hand, high YP can contribute to yield in moderate stress environments. Plant production in water-limited environments is very often affected by constitutive plant traits that allow maintenance of a high plant water status (dehydration avoidance). Osmotic adjustment (OA) is a major cellular stress adaptive response in certain crop plants that enhances dehydration avoidance and supports yield under stress. Despite past voiced speculations, there is no proof that OA entails a cost in terms of reduced YP. WUE for yield is often equated in a simplistic manner with DR. The large accumulation of knowledge on crop WUE as derived from research on carbon isotope discrimination allows some conclusions on the relations between WUE on the one hand, and DR and YP on the other, to be made. Briefly, apparent genotypic variations in WUE are normally expressed mainly due to variations in water use (WU; the denominator). Reduced WU, which is reflected in higher WUE, is generally achieved by plant traits and environmental responses that reduce YP. Improved WUE on the basis of reduced WU is expressed in improved yield under water-limited conditions only when there is need to balance crop water use against a limited and known soil moisture reserve. However, under most dryland situations where crops depend on unpredictable seasonal rainfall, the maximisation of soil moisture use is a crucial component of drought resistance (avoidance), which is generally expressed in lower WUE. It is concluded that the effect of a single 'drought adaptive' gene on crop performance in water-limited environments can be assessed only when the whole system is considered in terms of YP, DR, and WUE.

Drought resistance, water-use efficiency, and yield potential-are they compatible, dissonant, or mutually exclusive?

This publication opens with the inevitable introduction, moves on to the present traditional approach to breeding for yield stability, and then enumerates a detailed discussion of the physiological approach to breeding for resistance to specific stresses. Not all environmental stresses are covered, omitting those for which little can be said today on practical breeding solutions.

Plant Breeding For Stress Environments

http://eprints.icrisat.ac.in/824/1/FCR112_119-123_2009.pdf

Effective use of water (EUW) and not water-use efficiency (WUE) is the target of crop yield improvement under drought stress

Drought is a multidimensional stress affecting plants at various levels of their organization. The effect of and plant response to drought at the whole plant and crop level is most complex because it reflects the integration of stress effects and responses at all underlying levels of organization over space and time. This review discusses some of the major aspects of crop response to drought stress which are relevant for plant breeding. Emphasis is given to whole plant aspects which are too often disregarded when conclusions are drawn from molecular studies towards the genetic improvement of crop drought resistance. Topics discussed are seedling emergence and establishment, plant phenology, leaf area, water deficit and assimilation, osmotic adjustment, the root and the formation of yield. The discussion is concluded with the interpretation of crop adaptation to drought conditions in its agronomic sense. Conclusions are drawn regarding plant breeding for drought-prone conditions.

Crop responses to drought and the interpretation of adaptation

Osmotic adjustment (OA) and cellular compatible solute accumulation are widely recognized to have a role in plant adaptation to dehydration mainly through turgor maintenance and the protection of specific cellular functions by defined solutes. At the same time, there has been an ongoing trickle of skepticism in the literature about the role of OA in supporting crop yield under drought stress. Contrarian reviews argued that OA did not sustain turgor or that it served mainly for plant survival rather than productivity. This critical review examined 26 published studies where OA was compared with yield under drought stress in variable genotypes of 12 crops, namely, barley, wheat, maize, sorghum, chickpea, pea, pigeon pea, soybean, canola, mustard, castor bean and sunflower. Over all crops a positive and significant association between OA and yield under drought stress were found in 24 out of 26 cases. Considering that it is generally difficult to find a singular plant trait responsible for yield advantage of numerous crops under different drought stress conditions, this evidence is no less than remarkable as proof that OA sustains crop yield under drought stress.

https://www.onlinelibrary.wiley.com/doi/pdf/10.1111/pce.12800

Abraham Blum

Papers

Drought resistance, water-use efficiency, and yield potential-are they compatible, dissonant, or mutually exclusive?

Plant Breeding For Stress Environments

Effective use of water (EUW) and not water-use efficiency (WUE) is the target of crop yield improvement under drought stress

Crop responses to drought and the interpretation of adaptation

Osmotic adjustment is a prime drought stress adaptive engine in support of plant production