Breeding for low input conditions and consequences for participatory plant breeding: Examples from tropical maize and wheat

Participatory plant breeding (PPB) has been suggested as an effective alternative to formal plant breeding (FPB) as a breeding strategy for achieving productivity gains under low input conditions. With genetic progress through PPB and FPB being determined by the same genetic variables, the likelihood of success of PPB approaches applied in low input target conditions was analyzed using two case studies from FPB that have resulted in significant productivity gains under low input conditions: (1) breeding tropical maize for low input conditions by CIMMYT, and (2) breeding of spring wheat for the highly variable low input rainfed farming systems in Australia. In both cases, genetic improvement was an outcome of long-term investment in a sustained research effort aimed at understanding the detail of the important environmental constraints to productivity and the plant requirements for improved adaptation to the identified constraints, followed up by the design and continued evaluation of efficient breeding strategies. The breeding strategies used differed between the two case studies but were consistent in their attention to the key determinants of response to selection: (1) ensuring adequate sources of genetic variation and high selection pressures for the important traits at all stages of the breeding program, (2) use of experimental procedures to achieve high levels of heritability in the breeding trials, and (3) testing strategies that achieved a high genetic correlation between performance of germplasm in the breeding trials and under on-farm conditions. The implications of the outcomes from these FPB case studies for realizing the positive motivations for adopting PPB strategies are discussed with particular reference for low input target environment conditions.

Identification of causal relationships among traits related to drought resistance in Stylosanthes scabra using QTL analysis

Previous studies have shown that a negative relationship exists between transpiration efficiency (TE) and carbon isotope discrimination (Delta) and between TE and specific leaf area (SLA) in Stylosanthes scabra, A glasshouse experiment was conducted to confirm these relationships in an F-2 population and to study the causal nature of these relationships through quantitative trait loci (QTL) analysis, One hundred and twenty F-2 genotypes from a cross between two genotypes within S. scabra were used. Three replications for each genotype were maintained through vegetative propagation, Water stress was imposed by maintaining plants at 40% of field capacity for about 45 d. To facilitate QTL analysis, a genetic linkage map consisting of 151 RAPD markers was developed, Results from this study show that Delta was significantly and negatively correlated with TE and biomass production. Similarly, SLA showed significant negative correlation with TE and biomass production, Most of the QTL for TE and Delta were present on linkage groups 5 and 11. Similarly, QTL for SLA, transpiration and biomass productivity traits were clustered on linkage groups 13 and 24, One unlinked marker was also associated with these traits, There were several markers coincident between different traits, At all the coincident QTL, the direction of QTL effects was consistent with phenotypic data, At the coincident markers between TE and Delta, high alleles of TE were associated with low alleles of Delta. Similarly, low alleles of SLA were associated with high alleles of biomass productivity traits and transpiration. At the coincident markers between trans-4-hydroxy-N-methyl proline (MHP) and relative water content (RWC), low alleles of MHP were associated with high alleles of RWC, This study suggests the causal nature of the relationship between TE and Delta. Phenotypic data and QTL, data show that SLA was more closely associated with biomass production than with TE, This study also shows that a cause-effect relationship may exist between SLA and biomass production.

Yield response of rice (Oryza sativa L.) genotypes to different types of drought under rainfed lowlands - Part 3. Plant factors contributing to drought resistance

A series of experiments were conducted in drought-prone northeast Thailand to examine the magnitude of yield responses of diverse genotypes to drought stress environments and to identify traits that may confer drought resistance to rainfed lowland rice. One hundred and twenty eight genotypes were grown under non-stress and four different types of drought stress conditions. Under severe drought conditions, the maintenance of PWP of genotypes played a significant role in determining final grain yield. Because of their smaller plant size (lower total dry matter at anthesis) genotypes that extracted less soil water during the early stages of the drought period, tended to maintain higher PWP and had a higher fertile panicle percentage, filled grain percentage and final grain yield than other genotypes. PWP was correlated with delay in flowering (r = -0.387) indicating that the latter could be used as a measure of water potential under stress. Genotypes with well-developed root systems extracted water too rapidly and experienced severe water stress at flowering. RPR which showed smaller coefficient of variation was more useful than root mass density in identifying genotypes with large root system. Under less severe and prolonged drought conditions, genotypes that could achieve higher plant dry matter at anthesis were desirable. They had less delay in flowering, higher grain yield and higher drought response index, indicating the importance of ability to grow during the prolonged stress period. Other shoot characters (osmotic potential, leaf temperature, leaf rolling, leaf death) had little effect on grain yield under different drought conditions. This was associated with a lack of genetic variation and difficulty in estimating trait values precisely. Under mild stress conditions (yield loss less than 50%), there was no significant relationship between the measured drought characters and grain yield. Under these mild drought conditions, yield is determined more by yield potential and phenotype than by drought resistant mechanisms per se. (C) 2002 Elsevier Science B.V. All rights reserved.

An investigation into the possible inhibitory effects of white cypress pine (Callitris glaucophylla) litter on the germination and growth of associated ground cover species

White cypress-pine stands typically support sparse densities of shrubs and grasses. The commonly held opinion is that leaching of allelopathic chemical compounds from cypress-pine litter partly facilitates this exclusion. Germination and growth of cypress pine seedlings do not appear to be similarly affected. This study set out to determine whether cypress litter had a differential effect on germination and growth of cypress-pine seedlings and on associated ground-cover species. Glasshouse trials comparing seedling emergence under cypress- and artificial-litter layers were undertaken. Cypress-pine litter did not have an inhibitory effect on the germination or growth of ground-cover species. In most cases, seedling emergence was facilitated by the application of cypress-pine litter due to its ability to increase the water holding capacity of the underlying soil. Cypress litter did not promote growth of its own seedlings over its competitors except on coarse-textured soils where it provided an ameliorative function to water stress due to the soil's reduced water holding capacity. The inhibition of ground-cover species' germination and growth in pure cypress stands was suggested to be the result of high below-ground resource competition due to the pine's expansive root morphology.

Storage protein mobilization and thiol protease up-regulation by solid matrix priming in loblolly pine (Pinus taeda) seed embryos

Experiments were conducted to investigate physiological mechanisms of solid matrix priming (SMP) on germination enhancement of loblolly pine (Pinus taeda) seeds. During SMP, osmotic potential in the embryo decreased by 0.65 MPa, concentration of crystalloid proteins decreased to 62% and concentrations of buffer soluble proteins and free amino acids increased by 22% and by 166%, respectively. Observations under an electron microscope demonstrated protein bodies in the embryo were mobilized. Inhibitor analysis indicated thiol protease was the dominant enzyme among endopiptidases to degrade the reserved proteins. A fragment of thiol protease was cloned from the primed seed embryos and it has high identities to those thiol proteases responsive to water stress. RNA get blot analysis showed a 1.5 kb thiol protease gene was up-regulated by SMP. Treatment with E64, a thiol protease inhibitor, negated SMP effects on germination performance, water potentials and protein profiles. Based on the experimental results, reserve protein mobilization induced by SMP in the embryo before radicle emergence might be one of the mechanisms to enhance germination in loblolly pine seeds.

Impact of root and shoot temperature on bud dormancy and floral induction in lychee (Litchi chinensis Sonn.)

Potted lychee trees (cv. Tai so) with mature vegetative flushes were grown under three day/night temperature regimes known to induce floral (18/13degreesC), intermediate (23/18degreesC) and vegetative (28/23degreesC) shoot structures. Heating roots respective to shoots accelerated bud-break and shoot emergence, but reduced the level of floral initiation in emergent shoots. At 18/13degreesC, root temperatures of 20 and 25degreesC decreased the period of shoot dormancy from 9 weeks to 5 and 3 weeks, respectively. A root temperature of 20degreesC also increased the proportion of both leafy and stunted panicles to normal leafless panicles, and reduced the number of axillary panicles accompanying each terminal particle. A root temperature of 25degreesC produced only vegetative shoots. At 23/18degreesC, heating roots increased the proportion of vegetative shoots and partially emerged buds to leafy and stunted particles as well as accelerating bud-break. Cooling of roots in relation to the shoot resulted in non-emergence of buds at both 28/23 and 23/18degreesC. Bud-break did not occur until root cooling was terminated and root temperature returned to that of the shoot. At 23/18degreesC, subsequent emergent shoots had a greater proportion of leafy panicles relative to control trees. At 28/23degreesC, all emergent shoots remained vegetative. Lychee floral initiation is influenced by both root and shoot temperature. Root temperature has a direct effect on the length of the shoot dormancy period, with high temperatures reducing this period and the subsequent level of floral initiation. However, an extended period of dormancy in itself is not sufficient for floral initiation, with low shoot temperatures also a necessary prerequisite. (C) 2003 Elsevier B.V. All rights reserved.

A simple regional-scale model for forecasting sorghum yield across North-Eastern Australia

Sorghum is the main dryland summer crop in NE Australia and a number of agricultural businesses would benefit from an ability to forecast production likelihood at regional scale. In this study we sought to develop a simple agro-climatic modelling approach for predicting shire (statistical local area) sorghum yield. Actual shire yield data, available for the period 1983-1997 from the Australian Bureau of Statistics, were used to train the model. Shire yield was related to a water stress index (SI) that was derived from the agro-climatic model. The model involved a simple fallow and crop water balance that was driven by climate data available at recording stations within each shire. Parameters defining the soil water holding capacity, maximum number of sowings (MXNS) in any year, planting rainfall requirement, and critical period for stress during the crop cycle were optimised as part of the model fitting procedure. Cross-validated correlations (CVR) ranged from 0.5 to 0.9 at shire scale. When aggregated to regional and national scales, 78-84% of the annual variation in sorghum yield was explained. The model was used to examine trends in sorghum productivity and the approach to using it in an operational forecasting system was outlined. (c) 2005 Elsevier B.V. All rights reserved.

A simulation model of cereal-legume intercropping systems for semi-arid regions I. Model development

Cereal-legume intercropping plays an important role in subsistence food production in developing countries, especially in situations of limited water resources. Crop simulation can be used to assess risk for intercrop productivity over time and space. In this study, a simple model for intercropping was developed for cereal and legume growth and yield, under semi-arid conditions. The model is based on radiation interception and use, and incorporates a water stress factor. Total dry matter and yield are functions of photosynthetically active radiation (PAR), the fraction of radiation intercepted and radiation use efficiency (RUE). One of two PAR sub-models was used to estimate PAR from solar radiation; either PAR is 50% of solar radiation or the ratio of PAR to solar radiation (PAR/SR) is a function of the clearness index (K-T). The fraction of radiation intercepted was calculated either based on Beer's Law with crop extinction coefficients (K) from field experiments or from previous reports. RUE was calculated as a function of available soil water to a depth of 900 mm (ASW). Either the soil water balance method or the decay curve approach was used to determine ASW. Thus, two alternatives for each of three factors, i.e., PAR/SR, K and ASW, were considered, giving eight possible models (2 methods x 3 factors). The model calibration and validation were carried out with maize-bean intercropping systems using data collected in a semi-arid region (Bloemfontein, Free State, South Africa) during seven growing seasons (1996/1997-2002/2003). The combination of PAR estimated from the clearness index, a crop extinction coefficient from the field experiment and the decay curve model gave the most reasonable and acceptable result. The intercrop model developed in this study is simple, so this modelling approach can be employed to develop other cereal-legume intercrop models for semi-arid regions. (c) 2004 Elsevier B.V. All rights reserved.

Adaptations of strangler figs to life in the rainforest canopy

Figs are rainforest keystone species. Non-strangler figs establish on the forest floor; strangler figs establish epiphytically, followed by a dramatic transition from epiphyte to free-standing tree that kills its hosts. Free-standing figs display vigorous growth and resource demand suggesting that epiphytic strangler figs require special adaptations to deal with resource limitations imposed by the epiphytic environment. We studied epiphytic and free-standing strangler figs, and non-strangler figs in tropical rainforest and in cultivation, as well as strangler figs in controlled conditions. We investigated whether the transition from epiphyte to free-standing tree is characterised by morphological and physiological plasticity. Epiphyte substrate had higher levels of plant-available ammonium and phosphate, and similar levels of nitrate compared with rainforest soil, suggesting that N and P are initially not limiting resources. A relationship was found between taxonomic groups and plant N physiology; strangler figs, all members of subgenus Urostigma, had mostly low foliar nitrate assimilation rates whereas non-strangler figs, in subgenera Pharmacocycea, Sycidium, Sycomorus or Synoecia, had moderate to high rates. Nitrate is an energetically expensive N source, and low nitrate use may be an adaptation of strangler figs for conserving energy during epiphytic growth. Interestingly, significant amounts of nitrate were stored in fleshy taproot tubers of epiphytic stranglers. Supporting the concept of plasticity, leaves of epiphytic Ficus benjamina L. had lower N and C content per unit leaf area, lower stomatal density and 80% greater specific leaf area than leaves of conspecific free-standing trees. Similarly, glasshouse-grown stranglers strongly increased biomass allocation to roots under water limitation. Epiphytic and free-standing F. benjamina had similar average foliar delta C-13, but epiphytes had more extreme values; this indicates that both groups of plants use the C-3 pathway of CO2 fixation but that water availability is highly variable for epiphytes. We hypothesise that epiphytic figs use fleshy stem tubers to avoid water stress, and that nitrate acts as an osmotic compound in tubers. We conclude that strangler figs are a unique experimental system for studying the transition from rainforest epiphyte to tree, and the genetic and environmental triggers involved.

Use of drought response index for identification of drought tolerant genotypes in rainfed lowland rice

Drought is a major constraint for rice production in the rainfed lowlands in Southeast Asia and Eastern India. The breeding programs for tainted lowland rice in these regions focus on adaptation to a range of drought conditions. However, a method of selection of drought tolerant genotypes has not been established and is considered to be one of the constraints faced by rice breeders. Drought response index (DRI) is based on grain yield adjusted for variation in potential yield and flowering date, and has been used recently, but its consistency among drought environments and hence its usefulness is not certain. In order to establish a selection method and subsequently to identify donor parents for drought resistance breeding, a series of experiments with 15 contrasting genotypes was conducted under well-watered and managed drought conditions at two sites for 5 years in Cambodia. Water level in the field was recorded and used to estimate the relative water level (WLREL) around flowering as an index of the severity of water deficit at the time of flowering for each entry. This was used to determine if DRI or yield reduction was due to drought tolerance or related to the amount of available water at flowering, i.e. drought escape. Grain yield reduction due to drought ranged from 12 to 46%. The drought occurred mainly during the reproductive phase, while four experiments had water stress from the early vegetative stage. There was significant variation for water availability around flowering among the nine experiments and this was associated with variation in mean yield reduction. Genotypic variation in DRI was consistent among most experiments, and genotypic mean DRI ranged from -0.54 to 0.47 (LSD 5% = 0.47). Genotypic variation in DRI was not related to WLREL around flowering in the nine environments. It is concluded that selection for DRI under drought conditions would allow breeders to identify donor lines with high drought tolerance as an important component of breeding better adapted varieties for the rainfed lowlands; two genotypes were identified with high DRI and low yield reduction and were subsequently used in the breeding program in Cambodia. (c) 2006 Elsevier B.V. All rights reserved.

Grain yield components of pearl millet under optimum conditions can be used to identify germplasm with adaptation to and zones

There is evidence that high-tillering, small-panicled pearl millet landraces are better adapted to the severe, unpredictable drought stress of the and zones of NW India than are low-tillering, large-panicled modern varieties, which significantly outyield the landraces under favourable conditions. In this paper, we analyse the relationship of and zone adaptation with the expression, under optimum conditions, of yield components that determine either the potential sink size or the ability to realise this potential. The objective is to test whether selection under optimal conditions for yield components can identify germplasm with adaptation to and zones in NW India, as this could potentially improve the efficiency of pearl millet improvement programs targeting and zones. We use data from an evaluation of over 100 landraces from NW India, conducted for two seasons under both severely drought-stressed and favourable conditions in northwest and south India. Trial average grain yields ranged from 14 g m(-2) to 182 g m(-2). The landraces were grouped into clusters, based on their phenology and yield components as measured under well-watered conditions in south India. In environments without pre-flowering drought stress, tillering type had no effect on potential sink size, but low-tillering, large-panicled landraces yielded significantly more grain, as they were better able to realise their potential sink size. By contrast, in two low-yielding and zone environments which experienced pre-anthesis drought stress, low-fillering, large-panicled landraces yielded significantly less grain than high-tillering ones with comparable phenology, because of both a reduced potential sink size and a reduced ability to realise this potential. The results indicate that the high grain yield of low-tillering, large-panicled landraces under favourable conditions is due to improved partitioning, rather than resource capture. However, under severe stress with restricted assimilate supply, high-tillering, small-panicled landraces are better able to produce a reproductive sink than are large-panicled ones. Selection under optimum conditions for yield components representing a resource allocation pattern favouring high yield under severe drought stress, combined with a capability to increase grain yield if assimilates are available, was more effective than direct selection for grain yield in identifying germplasm adapted to and zones. Incorporating such selection in early generations of variety testing could reduce the reliance on random stress environments. This should improve the efficiency of millet breeding programs targeting and zones. (c) 2005 Elsevier B.V. All rights reserved.

Contribution of potential yield, drought tolerance and escape to adaptation of 15 rice varieties in rainfed lowlands in Cambodia

In Cambodia, grain yield in rainfed lowland rice is often affected by drought during late vegetative or reproductive stage. Several experiments were conducted to quantify the contribution of potential yield, drought tolerance and drought escape mechanisms to yield under water stress conditions. In total nine pairs of well irrigated and simulated drought (by draining water) experiments were conducted. Potential yield was obtained under irrigation. Grain yields and flowering dates were recorded in 15 varieties. Drought tolerance was quantified by using drought response index (DRI), which is grain yield under drought adjusted for potential yield and flowering date of the variety. Drought escape is expressed as days to flower under drought conditions. Mean yield reduction due to drought of nine experiments was 27 % (range 12-44). The relative contribution of yield potential, flowering date and DRI to observe yield under drought were evaluated by multiple regression for each experiment. Potential yield accounted for 54% (with a range of 10-80) of the variation in actual yield under drought. This was followed by DRI and flowering date with 34 (with a range of 0-60) and 12 (with a range of 0-30) of the contribution, respectively. It is concluded that selecting for drought tolerance as well as for high yield potential would be important in developing cultivars for rainfed lowlands in Cambodia. Although flowering dates are important for drought escape, it had a small contribution probably because drought developed slowly in these experiments in Cambodia.

Leukocyte subset responses during exercise under heat stress with carbohydrate or water intake

Purpose: This study investigated leukocyte subset responses to moderate-intensity exercise under heat stress, with water (W) or carbohydrate (CHO) drink ingestion. Methods: In repeated trials, 13 soldiers consumed either a W or CHO drink during 3 h of walking at 4.4 km center dot h(-1) with a 5% gradient (15 min rest per hour) under heat stress (35 C and 55% relative humidity). The soldiers wore combat uniforms and carried water bottles and dummy rifles and ammunition, altogether weighing about 11.5 +/- 1.0 kg. Results: Plasma glucose concentration was significantly higher with CHO than W ingestion during exercise (p < 0.01). There were no significant differences between W and CHO conditions in exercise performance, plasma cortisol concentration, heart rate, or core temperature. CHO ingestion significantly moderated the increases in leukocyte (83% in W, 28% in CHO; p < 0.001), monocyte (60% in W, 34% in CHO; p < 0.05), and granulocyte counts (120% in W, 30% in CHO; p < 0.001), but not in lymphocyte count (41% in W, 25% in CHO). Conclusions: The increases in leukocyte and subset counts during moderate-intensity exercise under heat stress may be comparable to those observed during intense exercise in cool conditions. The response of immune cell counts is blunted by CHO intake during moderate-intensity exercise in the heat, and may not occur through the cortisol pathway.