Association mapping and genetic dissection of drought-induced canopy temperature differences in rice.

Drought-stressed plants display reduced stomatal conductance, which results in increased leaf temperature by limiting transpiration. In this study, thermal imaging was used to quantify the differences in canopy temperature under drought in a rice diversity panel consisting of 293 indica accessions. The population was grown under paddy field conditions and drought stress was imposed for 2 weeks at flowering. The canopy temperature of the accessions during stress negatively correlated with grain yield (r= -0.48) and positively with plant height (r=0.56). Temperature values were used to perform a genome-wide association (GWA) analysis using a 45K single nucleotide polynmorphism (SNP) map. A quantitative trait locus (QTL) for canopy temperature under drought was detected on chromosome 3 and fine-mapped using a high-density imputed SNP map. The candidate genes underlying the QTL point towards differences in the regulation of guard cell solute intake for stomatal opening as the possible source of temperature variation. Genetic variation for the significant markers of the QTL was present only within the tall, low-yielding landraces adapted to drought-prone environments. The absence of variation in the shorter genotypes, which showed lower leaf temperature and higher grain yield, suggests that breeding for high grain yield in rice under paddy conditions has reduced genetic variation for stomatal response under drought.

Improving intercropping: a synthesis of research in agronomy, plant physiology and ecology.

Intercropping is a farming practice involving two or more crop species, or genotypes, growing together and coexisting for a time. On the fringes of modern intensive agriculture, intercropping is important in many subsistence or low-input/resource-limited agricultural systems. By allowing genuine yield gains without increased inputs, or greater stability of yield with decreased inputs, intercropping could be one route to delivering 'sustainable intensification'. We discuss how recent knowledge from agronomy, plant physiology and ecology can be combined with the aim of improving intercropping systems. Recent advances in agronomy and plant physiology include better understanding of the mechanisms of interactions between crop genotypes and species ­ for example, enhanced resource availability through niche complementarity. Ecological advances include better understanding of the context-dependency of interactions, the mechanisms behind disease and pest avoidance, the links between above- and below-ground systems, and the role of microtopographic variation in coexistence. This improved understanding can guide approaches for improving intercropping systems, including breeding crops for intercropping. Although such advances can help to improve intercropping systems, we suggest that other topics also need addressing. These include better assessment of the wider benefits of intercropping in terms of multiple ecosystem services, collaboration with agricultural engineering, and more effective interdisciplinary research.

A new physical interpretation of plant root capacitance.

Capacitance has been used as a non-destructive measure of root system size for 30 years. The equipment required is cheap and simple to apply in both field and laboratory. Good linear correlations have been reported between capacitance and root mass. A model by F. N. Dalton, predicting a linear relationship between these two variables, has become accepted widely. This model was tested for barley (Hordeum vulgare) grown hydroponically using treatments that included: raising roots out of solution, cutting roots at positions below the solution surface, and varying the distance between plant electrode and the solution surface. Although good linear correlations were found between capacitance and mass for whole root systems, when roots were raised out of solution capacitances were not linearly related to submerged root mass. Excision of roots in the solution had negligible effect on the measured capacitance. These latter observations conflict with Dalton's model. Capacitance correlated linearly with the sum of root cross-sectional areas at the solution surface and inversely with distance between plant electrode and solution surface. A new model for capacitance is proposed that is consistent with these observations.

Field Phenomics: Will It Enable Crop Improvement?

Field phenomics has been identified as a promising enabling technology to assist plant breeders with the development of improved cultivars for farmers. Yet, despite much investment, there are few examples demonstrating the application of phenomics within a plant breeding program. We review recent progress in field phenomics and highlight the importance of targeting breeders' needs, rather than perceived technology needs, through developing and enhancing partnerships between phenomics researchers and plant breeders.

New phenotyping methods for screening wheat and barley for beneficial responses to water deficit.

This review considers stomatal conductance as an indicator of genotypic differences in the growth response to water stress. The benefits of using stomatal conductance are compared with photosynthetic rate and other indicators of genetic variation in water stress tolerance, along with the use of modern phenomics technologies. Various treatments for screening for genetic diversity in response to water deficit in controlled environments are considered. There is no perfect medium: there are pitfalls in using soil in pots, and in using hydroponics with ionic and non-ionic osmotica. Use of mixed salts or NaCl is recommended over non-ionic osmotica. Developments in infrared thermography provide new and feasible screening methods for detecting genetic variation in the stomatal response to water deficit in controlled environments and in the field.

The impact of drought on leaf physiology of Quercus suber L. trees: comparison of an extreme drought event with chronic rainfall reduction.

Understanding the responses of cork oak (Quercus suber L.) to actual and predicted summer conditions is essential to determine the future sustainability of cork oak woodlands in Iberia. Thermal imaging may provide a rapid method for monitoring the extent of stress. The ecophysiology of cork trees was studied over three years. Three treatments were applied by means of rainfall capture and irrigation, with plots receiving 120%, 100%, or 80% of natural precipitation. Despite stomatal closure, detected using both thermal imaging and porometry, leaf water potential fell during the summer, most drastically during the third year of accumulative stress. The quantum efficiency (ΦPSII) and the maximum efficiency Fv' /FM' of photosystem II also fell more intensely over the third summer, while non-photochemical quenching (NPQ) increased. The reduced precipitation treatment sporadically further reduced leaf water potential, stomatal conductance (gs), IG (an index of gs derived from thermal imaging), ΦPSII, and Fv' /FM', and increased leaf temperature and NPQ. It is concluded that these are very resilient trees since they were only severely affected in the third year of severe drought (the third year registering 45% less rainfall than average), and removing 20% of rainfall had a limited impact..

What plant is that? Tests of automated image recognition apps for plant identification on plants from the British flora.

There has been a recent explosion in development of image recognition technology and its application to automated plant identification, so it is timely to consider its potential for field botany. Nine free apps or websites for automated plant identification and suitable for use on mobile phones or tablet computers in the field were tested on a disparate set of 38 images of plants or parts of plants chosen from the higher plant flora of Britain and Ireland. There were large differences in performance with the best apps identifying >50 % of samples tested to genus or better. Although the accuracy is good for some of the top-rated apps, for any quantitative biodiversity study or for ecological surveys, there remains a need for validation by experts or against conventional floras. Nevertheless, the better-performing apps should be of great value to beginners and amateurs and may usefully stimulate interest in plant identification and nature. Potential uses of automated image recognition plant identification apps are discussed and recommendations made for their future use.

Singlet oxygen quenching by phenylamides and their parent compounds.

This paper demonstrates for the first time that plant metabolites of the phenylamide type, conjugates of putrescine with hydroxycinnamic acids (p-coumaric, caffeic and ferulic), possess 1O2 quenching properties. Data were obtained confirming that their acidic parent compounds were also able to quench 1O2, as did polyamines (putrescine, spermidine and spermine), and that this ability depends on the number of amino groups. Potentiation of the 1O2 quenching ability of the conjugates relative to both parent components was established. The importance of polyamines and phenylamides in the plant non-enzymatic antioxidant defence at sites of intensive 1O2 generation, such as the photosynthetic centers, was suggested.

Assessing Drought Responses Using Thermal Infrared Imaging.

Canopy temperature, a surrogate for stomatal conductance, is shown to be a good indicator of plant water status and a potential tool for phenotyping and irrigation scheduling. Measurement of stomatal conductance and leaf temperature has traditionally been done by using porometers or gas exchange analyzers and fine-wire thermocouples attached to the leaves, which are labor intensive and point measurements. The advent of remote or proximal thermal sensing technologies has provided the potential for scaling up to leaves, plants, and canopies. Thermal cameras with a temperature resolution of <0.1 K now allow one to study the temperature variation within and between plants. This chapter discusses some applications of infrared thermography for assessing drought and other abiotic and biotic stress and outlines some of the main factors that need to be considered when applying this to the study of leaf or canopy temperature whether in controlled environments or in the field.

Coping with drought: stress and adaptive responses in potato and perspectives for improvement.

Potato (Solanum tuberosum L.) is often considered as a drought sensitive crop and its sustainable production is threatened due to frequent drought episodes. There has been much research aiming to understand the physiological, biochemical, and genetic basis of drought tolerance in potato as a basis for improving production under drought conditions. The complex phenotypic response of potato plants to drought is conditioned by the interactive effects of the plant's genotypic potential, developmental stage, and environment. Effective crop improvement for drought tolerance will require the pyramiding of many disparate characters, with different combinations being appropriate for different growing environments. An understanding of the interaction between below ground water uptake by the roots and above ground water loss from the shoot system is essential. The development of high throughput precision phenotyping platforms is providing an exciting new tool for precision screening, which, with the incorporation of innovative screening strategies, can aid the selection and pyramiding of drought-related genes appropriate for specific environments. Outcomes from genomics, proteomics, metabolomics, and bioengineering advances will undoubtedly compliment conventional breeding strategies and presents an alternative route toward development of drought tolerant potatoes. This review presents an overview of past research activity, highlighting recent advances with examples from other crops and suggesting future research directions.

Chilling requirement of Ribes cultivars.

It is usually thought that adequate winter chill is required for the full flowering of many temperate woody species. This paper investigates the sensitivity of blackcurrant bud burst and flowering to natural weather fluctuations in a temperate maritime climate, and compares a range of chill models that have been proposed for assessing the accumulation of winter chill. Bud break for four contrasting cultivars are compared in an exceptionally cold and in a mild winter in Eastern Scotland. The results confirm the importance of chilling at temperatures lower than 0°C and demonstrate that no single chilling function applies equally to all blackcurrant cultivars. There is a pressing need for further model development to take into account the relationship between chilling temperatures and warming temperatures occurring both during and after the chill accumulation period.

Infra-red thermography for high throughput field phenotyping in Solanum tuberosum.

The rapid development of genomic technology has made high throughput genotyping widely accessible but the associated high throughput phenotyping is now the major limiting factor in genetic analysis of traits. This paper evaluates the use of thermal imaging for the high throughput field phenotyping of Solanum tuberosum for differences in stomatal behaviour. A large multi-replicated trial of a potato mapping population was used to investigate the consistency in genotypic rankings across different trials and across measurements made at different times of day and on different days. The results confirmed a high degree of consistency between the genotypic rankings based on relative canopy temperature on different occasions. Genotype discrimination was enhanced both through normalising data by expressing genotype temperatures as differences from image means and through the enhanced replication obtained by using overlapping images. A Monte Carlo simulation approach was used to confirm the magnitude of genotypic differences that it is possible to discriminate. The results showed a clear negative association between canopy temperature and final tuber yield for this population, when grown under ample moisture supply. We have therefore established infrared thermography as an easy, rapid and non-destructive screening method for evaluating large population trials for genetic analysis. We also envisage this approach as having great potential for evaluating plant response to stress under field conditions.

Effect of salinity on water relations of wild barley plants differing in salt tolerance.

BACKGROUND AND AIMS: Certain lines of wild barley (Hordeum spontaneum) are more tolerant of salinity than others. The physiological basis of this difference is examined in a comparative study of a saline-tolerant and saline-intolerant line that emphasizes plant water relations. METHODOLOGY: Effects of salt-treatment (75 mM maximum) extending from a few hours to 3 weeks were quantified in 8-day-old seedlings of a saline-sensitive wild barley line ('T-1') and a less saline-sensitive line ('20-45'). Plants were grown in nutrient culture. Levels of mRNA of the HtPIP2;4 aquaporin (AQP) gene were determined together with a range of physiological responses including root hydraulic conductivity, osmotic potential of root xylem sap, transpiration, leaf relative water content, root water content, leaf water potential, leaf sap osmolality, leaf length, leaf area and chlorophyll content. PRINCIPAL RESULTS: Salt treatment inhibited transpiration and hydraulic conductivity more in salt-tolerant '20-45' plants than in salt-sensitive 'T-1'. In '20-45', the effect was paralleled by a fast (within a few hours) and persistent (3 days) down-regulation of aquaporin. In salt-sensitive 'T-1' plants, aquaporin down-regulation was delayed for up to 24 h. Greater tolerance in '20-45' plants was characterized by less inhibition of leaf area, root fresh weight, leaf water content and chlorophyll concentration. Leaf water potentials were similar in both lines. CONCLUSIONS: (i) Decline in hydraulic conductivity in salt-treated barley plants is important for stomatal closure, (ii) lowered transpiration rate is beneficial for salt tolerance, at least at the seedling stage and (iii) changes in AQP expression are implicated in the control of whole plant hydraulic conductivity and the regulation of shoot water relations.

Multi-sensor plant imaging: Towards the development of a stress-catalogue.

Agricultural production is limited by a wide range of abiotic (e.g. drought, waterlogging) and biotic (pests, diseases and weeds) stresses. The impact of these stresses can be minimized by appropriate management actions such as irrigation or chemical pesticide application. However, further optimization requires the ability to diagnose and quantify the different stresses at an early stage. Particularly valuable information of plant stress responses is provided by plant imaging, i.e. non-contact sensing with spatial resolving power: (i) thermal imaging, detecting changes in transpiration rate and (ii) fluorescence imaging monitoring alterations in photosynthesis and other physiological processes. These can be supplemented by conventional video imagery for study of growth. An efficient early warning system would need to discriminate between different stressors. Given the wide range of sensors, and the association of specific plant physiological responses with changes at particular wavelengths, this goal seems within reach. This is based on the organization of the individual sensor results in a matrix that identifies specific signatures for multiple stress types. In this report, we first review the diagnostic effectiveness of different individual imaging techniques and then extend this to the multi-sensor stress-identification approach.

Thermal infrared imaging of crop canopies for the remote diagnosis and quantification of plant responses to water stress in the field.

Thermal imaging using infrared (IR) is now an established technology for the study of stomatal responses and for phenotyping plants for differences in stomatal behaviour. This paper outlines the potential applications of IR sensing in drought phenotyping, with particular emphasis on a description of the problems with extrapolation of the technique from the study of single leaves in controlled environments to the study of plant canopies is field plots, with examples taken from studies on grapevine (Vitis vinifera L.) and rice (Oryza sativa L.). Particular problems include the sensitivity of leaf temperature (and potentially the temperature of reference surfaces) to both temporal and spatial variation in absorbed radiation, with leaf temperature varying by as much as 15°C between full sun and deep shade. Examples of application of the approach to phenotyping in the field and the steps in data analysis are outlined, demonstrating that clear genotypic variation may be detected despite substantial variation in soil moisture status or incident radiation by the use of appropriate normalisation techniques.

Monitoring plant and soil water status: established and novel methods revisited and their relevance to studies of drought tolerance.

In all studies of the effects of water deficits on plant functioning there is a need for an accurate and comprehensive definition of treatments and their effects on plant water status. The various measures of water status used in plant and soil science are reviewed and their appropriateness for different purposes such as for studies of mechanistic effects of water deficits on plants, for breeding of drought-tolerant plants, or for management of irrigation systems are reviewed. An important conclusion is that the frequent emphasis on water potential rather than on cell turgor can be shown to be misleading, as can be measurements in the leaf. The disadvantages of the current trend towards the omission of necessary water-status measurements, especially common in more molecular studies, are outlined, and recommendations made for minimal sets of measurements for specific types of experiments.

Response of photosynthetic apparatus to moderate high temperature in contrasting wheat cultivars at different oxygen concentrations.

The photosynthetic responses to moderately high temperatures (38 degrees C, imposed at 21% or 2% O(2) in air and 1500 mumol m(-2) s(-1)) were compared in wheat (Triticum aestivum L.) cultivars grown in northern regions of Ukraine and expected to be relatively sensitive to high temperatures ('North' cultivars) and in cultivars grown in southern regions and expected to be relatively heat-tolerant ('South' cultivars). Heating intact leaves in 21% O(2) for 1 h decreased CO(2) assimilation by c. 63% in 'North' cultivars and only c. 32% in 'South' cultivars, with a decrease in PSII activity being observed in only one of the 'North' cultivars. Carboxylation efficiency was decreased by about 2.7-fold in 'North' cultivars with no significant effect in 'South' cultivars. The maximum rates of carboxylation by Rubisco in vivo, V(cmax), estimated from Farquhar's model, increased more than 2-fold in 'South' cultivars and remained unchanged in 'North' cultivars while the maximum rate of RuBP regeneration, J(max), decreased by 53% and 21% in 'North' and 'South' cultivars, respectively. Where the heat treatment was imposed in 2% O(2) this increased (as compared with treatment at 21% O(2)) the inhibitory effect on CO(2) assimilation in tolerant cultivars, but decreased it in sensitive ones. The results suggested that differences in tolerance of moderately high temperatures in wheat relate to the stability of the Rubisco function and to RuBP regeneration activity rather than to the effects on PSII activity or stomatal control.