Putrescine Increases Floral Retention, Pod Set and Seed Yield in Cold Stressed Chickpea

Chickpea (Cicer arietinum L.) is sensitive to cold stress (<8 °C) at its reproductive phase that results in flower abortion, poor pod set and thus reduced yield. Early maturing genotypes are especially more sensitive. In this crop, the metabolic causes underlying cold injury that are imperative to induce cold tolerance are not known. In the present study, the endogenous levels of putrescine (diamine), spermidine (triamine) and spermine (tetramine) were examined in early maturing chickpea genotype ICCV 96029, subjected to chilling temperatures of field (12–15/4–6 °C; average maximum and minimum temperature respectively), at flowering or early podding stage. These were compared with controls growing in warmer conditions (28/12 °C) of the glasshouse. The polyamine levels increased six to nine times because of stress. Relatively, putrescine (PUT) elevation was the highest but short-lived and its decrease appeared to match with the onset of flower and pod abscission in stressed plants. Compared with controls, chilling injury, observed as electrolyte leakage (EL), increased by 60 % while cellular respiration declined by 68 % in stressed plants. Exogenous application of 10 mm PUT to stressed plants reduced the EL by 29 % and elevated the cellular respiration by 40 %. PUT application at flowering stage resulted in increase of 30, 31, 23 and 25 % in floral retention, pod set, pod retention and fertile pods respectively. At the early podding stage, PUT treatment increased the seed yield per plant, seed number per 100 pods and individual seed weight by 50, 17 and 19 % respectively. The number of single-seeded pods per plant increased from 4.4 in stressed plants to 12.2 in PUT-treated plants while the number of double-seeded pods reduced from 6.2 to 4.3. The number of infertile pods declined from 8.2 in stressed plants to 3.1 in PUT-treated plants.     

Heat stress in food legumes: evaluation of membrane thermostability methodology and use of infra-red thermometry

Heat tolerance in 45 chickpea, lentil, and faba bean genotypes was investigated during 2007/2008 and 2008/2009 at Alexandria Agriculture Research Station, Alexandria, Egypt, using screening methods employing the membrane thermostability technique. Threshold temperature to be used in screening for heat tolerance at germination was also investigated for each crop. Temperatures, responsible for 50% germination were 40, 33.5, and 29°C for chickpea, faba bean, and lentil, respectively. Germination percent under high temperature varied significantly (P ≤ 0.05) amongst genotypes. Germination percentage ranged from 4.8 to 71.6, 39.2 to 90.0, and 4.8 to 68.6, in chickpea, lentil, and faba bean, respectively. Differences were significant (P ≤ 0.05) among faba bean and chickpea genotypes. Membrane relative injury (RI%) showed significant (P ≤ 0.05) variability among the genotypes and ranged from 10.57 to 58, 5.2 to 61.7, and 15.7 to 52.7 in chickpea, lentil, and faba bean, respectively. Canopy temperature was measured to evaluate heat avoidance in tested genotypes. Infra-red thermometry was used to measure canopy temperature and the gradient of canopy to ambient air temperature (∆T_C-A) in moisture stressed and unstressed treatments. Canopy temperature, leaf water potential (LWP) and leaf water content were affected by the level of soil moisture. Genotypes were able to bring their canopy temperatures to levels lower than ambient air temperatures but the differences were not significant. A heat stress index (HSI) were computed relating the ∆T_C-A in moisture stressed to unstressed treatments. Regression of leaf water potential (LWP) and the heat stress index (HSI) was significant (P ≤ 0.05) in faba bean genotypes in the stressful environment. The results of the present investigation emphasize the efficiency of membrane thermostability technique in selection for heat tolerance in early stages of growth in food legumes.     

Screening techniques and sources of resistance to abiotic stresses in cool-season food legumes

Summary The adaptability and productivity of cool-season food legumes (chickpea, faba bean, lentil, pea) are limited by major abiotic stresses including drought, heat, frost, chilling, waterlogging, salinity and mineral toxicities. The severity of these stresses is unpredictable in field experiments, so field trials are increasingly supplemented with controlled-environment testing and physiological screening. For drought testing, irrigation is used in dry fields and rain-out shelters in damp ones. Carbon isotope discrimination (Δ¹³C) is a well-established screen for drought tolerance in C3 cereal crops which is now being validated for use in grain legumes, but it is relatively expensive per sample and more economical methods include stomatal conductance and canopy temperature. Chickpea lines ICC4958 and FLIP87-59C and faba bean line ILB938 have demonstrated good drought tolerance parameters in different experiments. For frost tolerance, an efficient controlled-environment procedure involves exposing hardened pot-grown plants to sub-zero temperatures. Faba beans Cote d’Or and BPL4628 as well as lentil ILL5865 have demonstrated good freezing tolerance in such tests. Chilling-tolerance tests are more commonly conducted in the field and lentil line ILL1878 as well as derivatives of interspecific crosses between chickpea and its wild relatives have repeatedly shown good results. The timing of chilling is particularly important as temperatures which are not lethal to the plant can greatly disrupt fertilization of flowers. Salinity response can be determined using hydroponic methods with a sand or gravel substrate and rapid, efficient scoring is based on leaf symptoms. Many lines of chickpea, faba bean and lentil have shown good salinity tolerance in a single article but none has become a benchmark. Waterlogging tolerance can be evaluated using paired hydroponic systems, one oxygenated and the other de-oxygenated. The development of lysigenous cavities or aerenchyma in roots, common in warm-season legumes, is reported in pea and lentil but is not well established in chickpea or faba bean. Many stresses are associated with oxidative damage leading to changes in chlorophyll fluorescence, membrane stability and peroxidase levels. An additional factor relevant to the legumes is the response of the symbiotic nitrogen-fixing bacteria to the stress.     

Soybean Pollen Anatomy,Viability and Pod Set under High Temperature Stress

High temperature (HT) stress is one of the major environmental factors influencing yield of soybean (Glycine max L. Merr.) in the semi‐arid regions. Experiments were conducted in controlled environments to study the effects of HT stress on anatomical changes of pollen and their relationship to pollen function in soybean genotype K 03‐2897. Objectives of this study were to (a) quantify the effect of HT stress during flowering on pollen function and pod set and (b) observe the anatomical changes in pollen grains of soybean plants grown under HT stress. Plants were exposed to HT (38/28 °C) or optimum temperature (OT, 28/18 °C) for 14 days at flowering stage. HT stress significantly decreased in vitro pollen germination by 22.7 % compared to OT. Pollen from HT stress was deformed; it had a thicker exine wall and a disintegrated tapetum layer. HT stress decreased pod set percentage (35.2 %) compared to OT. This study showed that decreases in pollen in vitro germination by HT stress were caused by anatomical changes in pollen, leading to decreased pod set percentage under HT stress.     

Exogenous Application of Abscisic Acid Improves Cold Tolerance in Chickpea (Cicer arietinum L.)

Chickpea suffers cold stress (<10 °C) damage especially during reproductive phase resulting in the abortion of flowers and pods, poor pod set, and reduction in seed yield and seed quality. One of the ways in modifying cold tolerance involves exogenous treatment of the plants with chemicals having established role in cold tolerance. In the present study, the chickpea plants growing under optimum temperature conditions (28/12 °C, as average maximum and minimum temperature) were subjected to cold conditions of the field (10–12/2–4 °C; day/night as average maximum and minimum temperature) at the bud stage. Prior to exposure, these plants were treated exogenously with 10 μm abscisic acid (ABA) and thereafter again after 1 week of exposure. The stress injury measured in terms of increase in electrolyte leakage, decrease in 2,3,5-triphenyl tetrazolium chloride reduction %, relative leaf water content and chlorophyll content was observed to be significantly mitigated in ABA-applied plants. A greater pollen viability, pollen germination, flower retention and pod set were noticed in ABA-treated plants compared with stressed plants. The seed yield showed considerable improvement in the plants treated with ABA relative to the stressed plants that was attributed to the increase in seed weight, greater number of single seeded pods and reduction in number of infertile pods. The oxidative damage measured as thiobarbituric acid-reactive substances was lesser in ABA-treated plants that was associated with greater activities of superoxide dismutase, catalase, ascorbate peroxidase, ascorbic acid, glutathione and proline in these plants. It was concluded that cold stress effects were partly overcome by ABA treatment because of the improvement in water status of the leaves as well as the reduction in oxidative damage.     

Temperature and relative humidity effects on quality,total ascorbic acid,phenolics and flavonoid concentrations,and antioxidant activity of strawberry

The physical qualities and antioxidant components of ‘Jewel’ strawberry fruit stored in 75, 85 or 95% relative humidity (RH) at 0.5, 10 and 20 °C for 4 days were studied. Overall fruit quality declined more rapidly at 20 °C, especially at 95% RH. Weight loss of fruit was negligible for 2 days at all temperatures but it increased at 10 °C in the lowest RH and increased rapidly from day 3 at 20 °C especially with lower RH. Firmness was maintained, or even increased, at 0.5 or 10 °C, while soluble solids concentrations (SSC) decreased at higher storage temperatures. Red color, assessed using chroma, hue and lightness, and anthocyanin concentrations were relatively unchanged at 0.5 or 10 °C but increased rapidly at 20 °C as fruit ripened. Firmness, SSC and color were not affected by RH. Total phenolic compounds were slightly higher at 20 °C than at other temperatures at all RHs. Total ascorbic acid concentrations of the fruit remained similar for the first 2 days of storage, then declined in fruit stored at 0.5 and 20 °C, but remained unchanged at 10 °C at all RHs. Total flavonoid content of fruit did not change over time at all temperatures. The total antioxidant activity of fruit was higher at 10 °C than at 0.5 and 20 °C on day 3, and no effect of RH was detected. In conclusion, while the best temperature for long-term storage is 0.5 °C, quality could be maintained at 10 °C for acceptable periods of time for marketing and may be associated with better nutritional quality.     

Identification and characterization of sources for photo‐ and thermo‐insensitivity in Vigna species

Cultivation of the same varieties of mungbean and blackgram across different seasons and locations is constrained by their photo‐ and thermo‐sensitive behaviour. Developing insensitive genotypes, which can fit well across all seasons, requires robust donors which would provide genes imparting this trait. This study was undertaken to identify such donors in the Vigna species. Forty‐eight accessions belonging to 13 Vigna species and eight released cultivars were evaluated under natural field conditions. Among these, two accessions, viz. V. umbellata (IC251442) and V. glabrescens (IC251372) were found photo‐ and thermo‐insensitive as these were able to flower and set pods at temperatures as high as 43.9°C and as low as 2.7°C. Pollen viability studies indicated viable pollen (>75% at 2.7°C and >85% at 41.9°C) and normal pollen tube growth at both the extremes of temperature. The identified V. glabrescens accession has long, constricted pods and dark green, mottled seeds while V. umbellata has smooth, curved pods and shining, oval, large seeds. Both these accessions can be utilized in developing photo–thermo insensitive genotypes in cultivated Vigna species.     

Screening chickpea for adaptation to water stress: Associations between yield and crop growth rate

Robust associations between yield and crop growth rate in a species-specific critical developmental window have been demonstrated in many crops. In this study we focus on genotype-driven variation in crop growth rate and its association with chickpea yield under drought. We measured crop growth rate using Normalised Difference Vegetative Index (NDVI) in 20 diverse chickpea lines, after calibration of NDVI against biomass accounting for morphological differences between Kabuli and Desi types. Crops were grown in eight environments resulting from the combination of seasons, sowing dates and water supply, returning a yield range from 152 to 366 g m⁻². For both sources of variation – environment and genotype – yield correlated with crop growth rate in the window 300 °Cd before flowering to 200 °Cd after flowering. In the range of crop growth rate from 0.07 to 0.91 g m⁻² °Cd⁻¹, the relationship was linear with zero intercept, as with other indeterminate grain legumes. Genotype-driven associations between yield and crop growth rate were stronger under water stress than under favourable conditions. Despite this general trend, lines were identified with high crop growth rate in both favourable and stress conditions. We demonstrate that calibrated NDVI is a rapid, inexpensive screening tool to capture a physiologically meaningful link between yield and crop growth rate in chickpea.     

Effect of Weed Removal on Productivity of Chickpea (Cicer arietinum L.) and Lentil (Lens culinaris Med.) in a Mediterranean Environment

Three field experiments were conducted on chickpea ( Cicer arietinum L.) and four on lentil ( Lens culinaris Med.) at different winter-sown rainfed locations in Jordan from 1988/89 to 1990/91 to study the effect of the duration of weed-free and weed-infested conditions on yields and yield components of the crops. Chickpea seed yields were reduced on average by 81 % and straw yields by 63 % when fields remained weed infested until harvest compared with weed-free conditions throughout the growing season. The corresponding lentil seed and straw yield decreases were 63 % and 55 %. As the duration of weed-free period increased and the duration of weed-infested period decreased, yields increased. However, the critical period of weed interference was between 35 and 49 days after emergence in chickpea and between 49 and 56 days after emergence in lentil, when these crops were at an advanced stage of vegetative growth. There were significant negative correlations between the weed dry weight and the seed or straw yields. The reduction in seed yields in both crops because of weed interference occurred mainly through the reduced number of pods /plant, which in turn was partly the result of reduced number of secondary branches. In chickpea, some reduction also occurred through reduced 100-seed weight.     

Impact of heat stress on pod-based yield components in field pea (Pisum sativum L.)

Elevated temperatures associated with climate change result in crops being exposed to frequent spells of heat stress. Heat stress results in reduced yield in field pea (Pisum sativum L.); it is therefore important to identify cultivars with improved pod and seed retention under heat to mitigate this loss. Objectives were to investigate the effect of heat stress on phenology, yield and pod-based yield components. Sixteen pea cultivars were evaluated at normal and late (hot) seeding dates in the field in Arizona 2012 and in growth chambers with two temperature regimes (24/18°C and 35/18°C day/night temperature for 7 days) during reproductive development. We measured variation in the pattern of pod retention at four-node positions on plants, seed retention by ovule position (stylar, medial and basal) within pods and screened cultivars for pod retention, seed retention and yield. Heat stress reduced seed yield by accelerating the crop lifecycle and reducing pod number and seed size. Heat stress had the most damaging effect on younger reproductive growth (flowers and pods developed later), resulting in ovary abortion from developing flowers. Heat also accelerated seed abortion in all ovule positions within pods. Two high-yielding cultivars under control temperature, “Naparnyk” and “CDC Meadow”, maintained high yield in heat, and “MFR043” had the lowest yield. Cultivars “40-10” and “Naparnyk” retained the most ovules and seeds per pod, and “MFR043” aborted seeds when exposed to heat. In half of the cultivars, ovules at the basal peduncle end of pods were likely to abort while ovules at the medial and stylar end positions developed into seeds. For seven of the field cultivars, ovules at the medial pod position also produced mature seeds. Cultivars “40-10”, “Naparnyk” and “CDC Meadow” had greater pod and ovule retention or maintained high yield under heat stress, and were identified as heat-tolerant cultivars. Our results allow for a better understanding of pod-based yield components in field pea under heat stress and developing heat-tolerant cultivars.     

Quality retention and potential shelf-life of fresh-cut lemons as affected by cut type and temperature

The effects of four cut types (wedges, slices, 1/2 and 1/4 slices) of ‘Lisbon’ lemons (Citrus lemon L.) and storage at four temperatures (0, 2, 5 and 10 °C) on post-cutting life were studied. Respiration rates of all cut types that were stored at 0, 2 and 5 °C up to 8 days were 2–5 times higher than those of the whole lemons, while the increase was up to 12-fold at 10 °C. Small differences among treatments were observed in the post-cutting changes of color parameters and chemical composition. Based on sensory analysis, the four cut types remained marketable for up to 7 days at all tested temperatures, but only the wedges, slices, and 1/2 slices stored at 0, 2 and 5 °C preserved their sensory attributes for up to 10 days. Good retention of vitamin C (about 85% ascorbic acid and 15% dehydroascorbic acid) and antioxidant capacity were found after 10 days at 0, 2, and 5 °C. Ethanol was the main fermentative metabolite found (88% of the total) and its concentration increased by up to three-fold in slices, 1/2 and 1/4 slices after 10 days at 10 °C. Total phenolics concentrations decreased gradually throughout the storage period in all cases.     

Postharvest conditioning of Satsuma mandarins for reduction of acidity and skin puffiness

Postharvest temperature and relative humidity (RH) treatments were tested for their capacity to increase the soluble solids content:titratable acidity ratio (SSC:TA ratio) and/or reduce skin puffiness of New Zealand grown ‘Miho’ Satsuma mandarin. Fruit of low SSC:TA (approximately 6.8:1) harvested in 2001 and 2002 were held at 18 or 30 °C at low (approximately 65%) or high (>95%) RH for 3 or 5 days, followed by 2 days at 10 °C (88–92% RH). In 2002, an additional treatment of high and low RH at 10 °C was examined. Treatments at 30 °C, irrespective of RH, resulted in increased SSC:TA ratios in the fruit as a result of a decrease in titratable acidity, largely a decrease in citric acid. There was little effect of temperature on SSC and the levels of individual sugars. There was no significant effect of RH on either TA or SSC. The altered metabolism was also seen in an elevated respiratory CO₂ output at 30 °C, but a decreased CO₂ output once these fruit were transferred to 10 °C, in comparison with fruit treated at lower temperatures.At 30 °C, weight loss was up to 8.5% after 5 days under low RH, but <4% under high RH. Fruit with >4% weight loss tended to have an unacceptable level of dehydration of the skin. After 5 days at 30 °C and low RH, skin puffiness, quantified from magnetic resonance images taken before and after treatment, was reduced, although fruit tended to have soft skin that could be more prone to damage.It is concluded that short high temperature treatments such as 3–5 days at 30 °C can significantly raise the SSC:TA ratio in Satsuma mandarin through a reduction in TA, and conducting these treatments under a RH >90% minimises the risk of excessive weight loss and softening of the skin.     

Heat tolerance in food legumes as evaluated by cell membrane thermostability and chlorophyll fluorescence techniques

Summary The genotypic variation for heat tolerance in chickpea, groundnut, pigeonpea, and soya bean was evaluated by testing membrane stability and photosystem (PSII) function in leaves at high temperatures. The legumes could be ranked from heat-tolerant to sensitive in the order: groundnut, soya bean, pigeonpea and chickpea. The damage to cell membranes (as reflected by an increased leakage of electrolytes) and PS II (as reflected by a decrease in the ratio of variable to maximum fluorescence) was less, and recovery from heat stress was faster in groundnut than in other crops. Prior exposure of plants to 35°C for 24h led to a reduced leakage of electrolytes at high temperatures in all crops but the differences among legumes were consistent. Substantial genotypic variation for heat tolerance was found in all legumes. Membrane injury was negatively associated with specific leaf weight in groundnut (r=–0.69^**) and soya bean (r=–0.56^**) but not in the pulses. Electrolyte leakage and fluorescence ratio were negatively correlated in all legumes. The potential use of electrolyte leakage and fluorescence tests as screening procedures for breeding heat-tolerant legumes is discussed.Abbreviations RI relative injury - Fo initial fluorescence - Fm maximum fluorescence - Fv variable fluorescence - PS II photosystem II - PAR photosynthetically active radiation     

Chickpea evolution has selected for contrasting phenological mechanisms among different habitats

Arguably the most important adaptive criterion in annual crops is appropriate phenology that minimizes exposure to climatic stresses and maximizes productivity in target environments. To date this has been achieved empirically by selecting among diverse genotypes in target locations. This approach is likely to become inadequate with pending climate change because selection is imposed on the outcome (flowering time) rather than the underlying mechanism (i.e. responses to daylength, ambient or vernalizing temperatures). In contrast to the cereals, in legumes the interaction between phenological mechanisms and environmental selection pressure is largely unknown. This paper addresses this shortcoming through photothermal modelling of chickpea germplasm from the world’s key production areas using a meta-analysis of multi-environment trials located from 49° N to 35° S. Germplasm origin had significant effects on temperature and daylength responsiveness, the former strongly correlated to vegetative phase temperatures at the collection or development site (r = 0.8). Accordingly, temperature responses increase from winter- to spring-sown Mediterranean and Australian material, and then to north, central & southern India. Germplasm origin also affects the relationship between photoperiod and temperature response. In Eastern Mediterranean material a strong negative relationship (r = −0.77) enables temperature insensitive genotypes to compensate through a strong photoperiod response. Clearly, chickpea evolution has selected for different phenological mechanisms across the habitat range. Given that under the anticipated global warming temperature sensitive cultivars will flower relatively earlier than those responding largely to photoperiod, it is important to exploit this diversity in developing better-adapted genotypes for future cropping environments.     

Capturing genetic variability and selection of traits for heat tolerance in a chickpea recombinant inbred line (RIL) population under field conditions

Chickpea is the most important pulse crop globally after dry beans. Climate change and increased cropping intensity are forcing chickpea cultivation to relatively higher temperature environments. To assess the genetic variability and identify heat responsive traits, a set of 296 F_8–9 recombinant inbred lines (RILs) of the cross ICC 4567 (heat sensitive) × ICC 15614 (heat tolerant) was evaluated under field conditions at ICRISAT, Patancheru, India. The experiment was conducted in an alpha lattice design with three replications during the summer seasons of 2013 and 2014 (heat stress environments, average temperature 35 °C and above), and post-rainy season of 2013 (non-stress environment, max. temperature below 30 °C). A two-fold variation for number of filled pods (FPod), total number of seeds (TS), harvest index (HI), percent pod setting (%PodSet) and grain yield (GY) was observed in the RILs under stress environments compared to non-stress environment. A yield penalty ranging from 22.26% (summer 2013) to 33.30% (summer 2014) was recorded in stress environments. Seed mass measured as 100-seed weight (HSW) was the least affected (6 and 7% reduction) trait, while %PodSet was the most affected (45.86 and 44.31% reduction) trait by high temperatures. Mixed model analysis of variance revealed a high genotypic coefficient of variation (GCV) (23.29–30.22%), phenotypic coefficient of variation (PCV) (25.69–32.44%) along with high heritability (80.89–86.89%) for FPod, TS, %PodSet and GY across the heat stress environments. Correlation studies (r = 0.61–0.97) and principal component analysis (PCA) revealed a strong positive association among the traits GY, FPod, VS and %PodSet under stress environments. Path analysis results showed that TS was the major direct and FPod was the major indirect contributors to GY under heat stress environments. Therefore, the traits that are good indicators of high grain yield under heat stress can be used in indirect selection for developing heat tolerant chickpea cultivars. Moreover, the presence of large genetic variation for heat tolerance in the population may provide an opportunity to use the RILs in future-heat tolerance breeding programme in chickpea.     

Mapping genes for double podding and other morphological traits in chickpea

Seed traits are important considerations for improving yield and product quality of chickpea (Cicer arietinum L.). The purpose of this study was to construct an intraspecific genetic linkage map and determine map positions of genes that confer double podding and seed traits using a population of 76 F₁₀ derived recombinant inbred lines (RILs) from the cross of ‘ICCV-2’ (large seeds and single pods) × ‘JG-62’ (small seeds and double podded). We used 55 sequence-tagged microsatellite sites (STMS), 20 random amplified polymorphic DNAs (RAPDs), 3inter-simple sequence repeats (ISSR) and 2 phenotypic markers to develop a genetic map that comprised 14 linkage groups covering297.5 cM. The gene for double podding (s) was mapped to linkage group 6 and linked to Tr44 and Tr35 at a distance of7.8 cM and 11.5 cM, respectively. The major gene for pigmentation, C, was mapped to linkage group 8 and was loosely linked to Tr33 at a distance of 13.5 cM. Four QTLs for 100 seed weight (located on LG4 and LG9), seed number plant^-1 (LG4), days to 50% flower (LG3) were identified. This intraspecific map of cultivated chickpea is the first that includes genes for important morphological traits. Synteny relationships among STMS markers appeared to be conserved on six linkage groups when our map was compared to the interspecific map presented by Winter et al. (2000). This revised version was published online in August 2006 with corrections to the Cover Date.     

Transfer of the Kosena Rfk1 gene,required in hybrid seed production,from oilseed rape to turnip rape

Radiant frost is a major abiotic stress, and one of the principal limiting factors for agricultural production worldwide, including Australia. Legumes, including field pea, faba bean, lentil and chickpea, are very sensitive to chilling and freezing temperatures, particularly at the flowering, early pod formation and seed filling stages. Radiant frost events occur when plants and soil absorb the sunlight during the day time and radiate heat during the night when the sky is clear and the air is still. Dense chilled air settles into the lowest areas of the canopy, where the most serious frost damage occurs. The cold air causes nucleation of the intracellular fluid in plant tissues and the subsequent rupturing of the plasma membrane. Among the cool season grain legume crops, chickpea, lentil and faba bean and field pea are the most susceptible to radiant frost injury during the reproductive stages. The more sensitive stages are flowering and podding. Frost at the reproductive stage results in flower abortion, poor pod set and impaired pod filling, leading to a drastic reduction in yield and quality. In contrast, in the UK and European countries, frost stress is related to the vegetative stages and, in particular, the effects of frost have been studied on cotyledon, uni/tri-foliolate leaf and seedling stages during the first few weeks of growth. Few winter genotypes have been identified as frost tolerant at vegetative stages. Vegetative frost tolerance is not related to reproductive frost tolerance, and hybrids from the vegetative frost-tolerant genotypes may not necessarily be tolerant at the reproductive stage. Tolerance to radiant frost has an inverse relationship with plant age. In the field, frost tolerance decreases from the vegetative stage to reproductive stage. Unlike wheat and barley, it is difficult to analyse and score frost damage in grain legume crops due to the presence of various phenophases on one plant at the reproductive stage. The extent of frost damage depends on the specific phenophases on a particular plant. However, current studies on genetic transformation of cold tolerant gene(s), membrane modifications, anti-freeze substances and ice nucleating or inhibiting agents provide useful information to improve our current understanding on frost damage and related mechanisms. The effects of frost damage on yield and grain quality illustrate the significance of this area of research. This review discusses the problem of radiant frost damage to cool season legumes in Australia and the associated research that has been carried out to combat this problem locally and worldwide. The available literature varies between species, specific climatic conditions and origin.     

Effect of abscisic acid applications on cold tolerance in chickpea (Cicer arietinum L.)

A series of field experiments were undertaken at three locations in Khyber PukhtunKhwa (KPK) Province, Pakistan to assess the effects of low temperatures and phytohormone applications on chickpea (Cicer arietinum L.) growth and yield. These trials showed that ABA application (10⁻⁴ M) to 40-day-old plants (before the first seasonal frost) offset low temperature-induced growth and yield depression at harvest (200-day-old plants) by up to 17%. These yield improvements were mainly due to an increase in the number of seeds pod⁻¹. Growth room experiments were carried out under controlled environmental conditions to establish how foliar application of 10⁻⁴ M ABA to 40-day-old plants might improve seed production at harvest. The foliar application of 10⁻⁴ M ABA had no detectable effect on endogenous shoot or root ABA levels four-days after spraying or on biomass when plants were maintained in warm conditions. When exposed to night temperatures of −2 °C, however, the endogenous ABA levels increased dramatically in both control and ABA-treated plants, but this rise was more rapid after ABA application (p < 0.01); after 14 days, these plants had gained significantly more biomass than the unsprayed controls (p < 0.05). No evidence was found to suggest ABA affected the osmotic or water balance of plants, but parallel experiments have shown ABA reduced low temperature-induced cell damage. Analysis of the proteome of the shoot tissues of ABA treated and untreated plants by 2-Dimensional Gel Electrophoresis identified several proteins that are induced by low temperatures and/or by ABA application in chickpea and which may be involved in conferring cold tolerance. Attempts were made to establish the identity of these proteins using mass spectrometry but in all cases the results were ambiguous; a more complete protein data base for legumes is required before the function of these proteins can be inferred.     

The Impact of Temperature on Seedling Root Traits of European and Tropical Maize (Zea mays L.) Cultivars

Twelve European and twelve tropical maize cultivars were grown in polyethylene tubes under three temperature regimes (14/12 °C, 22/20 °C; 30/28 °C). The plants were harvested when the third leaf was fully expanded. The experiments were undertaken in order to: (i) study the general response of root traits of maize seedlings to chilly, temperate and very warm conditions and (ii) find out whether and how the geographical origin of the maize material modifies the results. The results may be summarized as follows: 1. The 22/20 °C temperature regime led to the highest shoot and root dry weight, the greatest total root length, the most apices and the greatest root surface area. However, the optimal temperature for these traits appeared to be either slightly higher or lower than 22/20 °C (optimal temperature for number of apices > shoot dry weight > root surface area > root dry weight). 2. The shoot: root dry weight ratio was almost the same at 14/12 °C and 22/20 °C but increased significantly at 30/28 °C. The root surface area: root dry weight ratio was optimal at 22/20 °C but extremely low at 14/12 °C. The production of apices per unit root dry matter was lowest at 14/12 °C and increased steadily at higher temperatures. 3. The root surface area: shoot dry weight ratio was low both at 14/12 °C and at 30/28 °C. This may indicate that mineral nutrient deficiency, as a result of undersized root systems, is most likely at extremely low and extremely high temperatures. 4. Low temperatures promoted the formation of seminal roots. 5. Temperature affected the dominance of the longest seminal root (= primary root). At 14/12 °C, the primary root was rather short as compared to the seminal roots of higher order. At 30/28 °C, however, the primary root was markedly longer than other seminal roots of higher order. 6. At 14/12 °C, the European cultivars produced more shoot and root dry matter, generated more apices and developed greater root surface area than the tropical cultivars. At 30/28 °C, the situation was reversed. This confirms that the response to different temperatures is modified by the geographical origin of the plant material.