Brassinosteroids and the Tolerance of Cereals to Low and High Temperature Stress: Photosynthesis and the Physicochemical Properties of Cell Membranes

Cereals, which belong to the Poaceae family, are the most economically important group of plants. Among abiotic stresses, temperature stresses are a serious and at the same time unpredictable problem for plant production. Both frost (in the case of winter cereals) and high temperatures in summer (especially combined with a water deficit in the soil) can result in significant yield losses. Plants have developed various adaptive mechanisms that have enabled them to survive periods of extreme temperatures. The processes of acclimation to low and high temperatures are controlled, among others, by phytohormones. The current review is devoted to the role of brassinosteroids (BR) in cereal acclimation to temperature stress with special attention being paid to the impact of BR on photosynthesis and the membrane properties. In cereals, the exogenous application of BR increases frost tolerance (winter rye, winter wheat), tolerance to cold (maize) and tolerance to a high temperature (rice). Disturbances in BR biosynthesis and signaling are accompanied by a decrease in frost tolerance but unexpectedly an improvement of tolerance to high temperature (barley). BR exogenous treatment increases the efficiency of the photosynthetic light reactions under various temperature conditions (winter rye, barley, rice), but interestingly, BR mutants with disturbances in BR biosynthesis are also characterized by an increased efficiency of PSII (barley). BR regulate the sugar metabolism including an increase in the sugar content, which is of key importance for acclimation, especially to low temperatures (winter rye, barley, maize). BR either participate in the temperature-dependent regulation of fatty acid biosynthesis or control the processes that are responsible for the transport or incorporation of the fatty acids into the membranes, which influences membrane fluidity (and subsequently the tolerance to high/low temperatures) (barley). BR may be one of the players, along with gibberellins or ABA, in acquiring tolerance to temperature stress in cereals (particularly important for the acclimation of cereals to low temperature).

Mortality of Eucalyptus pilularis progenies and provenances after frost

We aimed to quantify frost tolerance and damage on Eucalyptus pilularis provenances and progenies in Curitibanos, Santa Catarina State, Brazil. Mortality and severity of damage were assessed before frost at 7 months old and at 14 and 19 months old after frost events. Frost resulted in plant mortalityof 55.7%, with no difference among progenies and provenances. The tolerance of 5.3% may have occurred due to specific local conditions (escape). We concluded that the species does not have tolerance to frost in subtropical environment, with daily thermal range from - 5 °C to 28 °C.

Sensitivity of Some Sweet Cherry (Prunus Avium L.) Genotypes To Late Spring Frosts During Different Phenological Stages Following Bud Burst

Sweet cherry (Prunus avium L.) is one of the economically important fruit crops worldwide. However, late spring frosts occurring in some years can significantly impact sweet cherry productivity through organ and tissue destruction caused by frost damage, and very little is known about frost tolerance or susceptibility of new cultivars. Differential thermal analysis (DTA) was, therefore, used to examine the exothermic characteristics (temperatures at which 50% of the flower buds were killed - mLTE values) of the flower buds belonging to members of the genus Prunus - 6 Prunus avium cultivars ('Noir de Guben', 'Bigarreau Gaucher', 'Merton Late', 'Merton Bigarreau', 'Van' and wild genotype). In the study, mLTE values of flowers of six cherry cultivars were determined at different floral bud developmental stages such as side green, green tip, open cluster, first white and full bloom under laboratory-based freeze assays for consecutive two years. The mLTE values of flower buds changed according to both different floral bud developmental stages and sweet cherry cultivars. In our findings, the mLTE values of flower buds in all cultivars generally occurred at higher temperatures in the open cluster stage, whereas the mLTE values of the flower buds in the first white stage occurred at lower temperatures, and therefore, these results are not only controversial in terms of previous frost tolerance studies, but also the first findings to be reported in literature. Considering the two-year average, the temperatures causing mLTE values for flower buds was -1.58 to -3.74°C at the side green stage, -0.94 to -3.51°C at the green tip stage, -0.41 to -1.96°C at the open cluster stage, -2.30 to -11.52°C at the first white stage and -2.37 to -9.80°C at the full bloom stage in the range of six cultivars. In laboratory-based freezing experiments, the 'Van' cultivar were least affected by low temperatures, followed by 'Bigarreau Gaucher' cultivar and wild genotype. 'Merton Late' cultivar, on the contrary, was the most sensitive cultivar to low temperatures, followed by 'Noir de Guben' and 'Merton Bigarreau'. These results can be valuable in predicting possible frost damage at different developmental stages of the flower buds in sweet cherry.

Extensive allele mining discovers novel genetic diversity in the loci controlling frost tolerance in barley

Key message Exome sequencing-based allele mining for frost tolerance suggests HvCBF14 rather than CNV at Fr-H2 locus is the main responsible of frost tolerance in barley. Abstract Wild relatives, landraces and old cultivars of barley represent a reservoir of untapped and potentially important genes for crop improvement, and the recent sequencing technologies provide the opportunity to mine the existing genetic diversity and to identify new genes/alleles for the traits of interest. In the present study, we use frost tolerance and vernalization requirement as case studies to demonstrate the power of allele mining carried out on exome sequencing data generated from > 400 barley accessions. New deletions in the first intron of VRN-H1 were identified and linked to a reduced vernalization requirement, while the allelic diversity of HvCBF2a, HvCBF4b and HvCBF14 was investigated by combining the analysis of SNPs and read counts. This approach has proven very effective to identify gene paralogs and copy number variants of HvCBF2 and the HvCBF4b-HvCBF2a segment. A multiple linear regression model which considers allelic variation at these genes suggests a major involvement of HvCBF14, rather than copy number variation of HvCBF4b-HvCBF2a, in controlling frost tolerance in barley. Overall, the present study provides powerful resource and tools to discover novel alleles at relevant genes in barley.

Genome Wide Association Study of Frost Tolerance in Wheat

Winter wheat growing areas in the Northern hemisphere are regularly exposed to heavy frost. Due to the negative impact on yield, the identification of genetic factors controlling frost tolerance (FroT) and development of tools for breeding is of prime importance. Here, we detected QTL associated with FroT by genome wide association studies (GWAS) using a diverse panel of 276 winter wheat genotypes that was phenotyped at five locations in Germany and Russia in three years. The panel was genotyped using the 90K iSelect array and SNPs in FroT candidate genes. In total, 17,566 SNPs were used for GWAS resulting in the identification of 53 markers significantly associated (LOD ≥4) to FroT, corresponding to 23 QTL regions located on 11 chromosomes (1A, 1B, 2A, 2B, 2D, 3A, 3D, 4A, 5A, 5B and 7D). The strongest QTL effect confirmed the importance of chromosome 5A for FroT. In addition, to our best knowledge, seven FroT QTLs were discovered for the first time in this study comprising QTLs on chromosomes 3A, 4A, 1B, and two on chromosomes 2D, 3D, and 7D. Identification of novel FroT candidate genes will help to better understand the FroT mechanism in wheat and to develop more effective combating strategies.

Study of the influence of negative temperatures on biomass accumulation and cell viability of callus and cell aggregates of Rhodiola rosea L.

As a result of determining the resistance to the action of different negative temperatures of callus cells and cellular aggregates of Rhodiola rosea, it was shown that after exposure of callus to -8oC, only 52% of the cells survived. In the case of exposing the experimental variant of R. rosea cell aggregates to -8oC, the value of cell viability was 68%. This suggests that the frost tolerance of cell aggregates is higher than that of callus cells, which indicates that the stress factor to be tolerated is higher, the lower the de-gree of organization of the biological system.

Genetic Variation in Frost Tolerance, Uromycladium acaciae Rust Resistance, and Growth in an Acacia mearnsii Population

Genetic variation in frost tolerance, resistance to the rust fungus Uromycladium acaciae, growth, stem form, and gummosis were evaluated in 110 open-pollinated families of black wattle (Acacia mearnsii De Wild). Families were tested at six frost-prone sites in northern KwaZulu-Natal and southeastern Mpumalanga, South Africa. Frost-hardy provenances were susceptible to rust disease and had poor growth. Locally grown F1 seed sources that originated from cold-hardy Australian seed sources had better growth and were tolerant to rust, but at a cost of lower frost tolerance. Considerable genetic variation was observed between families within seed sources for frost damage (hfm2 = 0.77), rust incidence (hfm2 = 0.89), and height (hfm2 = 0.80). The corresponding narrow-sense heritabilities (hi2) were 0.30, 0.80, and 0.32. Genotype-by-environment interaction levels were low for most traits. Except for a strong genetic correlation between tree height and diameter (0.90), all the pairs of traits had weak to moderate genetic correlations. Recurrent selection will be successful in improving frost tolerance and rust incidence. However, the current population comprises limited germplasm that is both tolerant to frost and resistant to rust. Thus, we recommend infusing germplasm from known cold-hardy Australian provenances into the current population to increase genetic variation for frost tolerance, rust, and growth. Study Implications Black wattle (Acacia mearnsii De Wild) is widely planted in South Africa for bark extract and woodchip exports. The species is prone to frost damage and susceptible to rust fungus Uromycladium acaciae, two major limitations to its cultivation. In this study, 110 open-pollinated families of black wattle originated from its native range in Australia and local South African seed sources were studied for frost and disease tolerance. The results from this study suggest that there is considerable variation within species for frost tolerance and fungal disease. We suggest a tandem selection strategy for black wattle farming in South Africa to increase resistance to frost damage and fungal diseases.

Methods for Measuring Frost Tolerance of Conifers: A Systematic Map

Frost tolerance is the ability of plants to withstand freezing temperatures without unrecoverable damage. Measuring frost tolerance involves various steps, each of which will vary depending on the objectives of the study. This systematic map takes an overall view of the literature that uses frost tolerance measuring techniques in gymnosperms, focusing mainly on conifers. Many different techniques have been used for testing, and there has been little change in methodology since 2000. The gold standard remains the field observation study, which, due to its cost, is frequently substituted by other techniques. Closed enclosure freezing tests (all non-field freezing tests) are done using various types of equipment for inducing artificial freezing. An examination of the literature indicates that several factors have to be controlled in order to measure frost tolerance in a manner similar to observation in a field study. Equipment that allows controlling the freezing rate, frost exposure time and thawing rate would obtain results closer to field studies. Other important factors in study design are the number of test temperatures used, the range of temperatures selected and the decrements between the temperatures, which should be selected based on expected frost tolerance of the tissue and species.

Seasonal Metabolic Investigation in Pomegranate (Punica granatum L.) Highlights the Role of Amino Acids in Genotype- and Organ-Specific Adaptive Responses to Freezing Stress

Every winter, temperate woody plants have to cope with freezing stress. Winter hardiness is of crucial importance for pomegranate survival and productivity. A comparative morphological and metabolic study was conducted on the stems and buds of 15 field-grown mature pomegranate genotypes in seven time-points during two developmental cycles. Seasonal changes of frost hardiness, as determined by electrolyte leakage method, and metabolite analysis by HPLC and GC revealed the variability in frost hardiness and metabolic contents result from genetic background and organ, as well as seasonal condition. Morphological adaptations, as well as metabolic remodeling, are the distinct features of the hardy genotypes. Larger buds with a greater number of compressed scales and the higher number of protective leaves, together with the higher number and content of changed metabolites, especially amino acids, seem to provide a higher frost resistance for those trees. We recorded two-times the change in metabolites and several-times accumulation of amino acids in the stem compared with buds. A better potential of stem for metabolome adjustment during the hardening period and a higher level of tolerance to stress is therefore suggested. High levels of arginine, proline, glutamine, and asparagine, and particularly the accumulation of alanine, tryptophan, and histidine are responsible for excellent tolerance of the stem of tolerant genotypes. With regard to the protective roles of amino acids, a relation between stress tolerance and the level of amino acids is proposed. This points both to the importance of amino acids in the winter survival of pomegranate trees, and to the evaluation of frost tolerance in other plants, by these specific markers.