Adaptive strategies to drought stress in grasses of the poaceae family under climate change: Physiological, genetic and molecular perspectives: A review.

Drought stress is one of the most critical abiotic factors which negatively impacts on growth, productivity, and survival of plants. Grass species have an important role in the sustainable intensification of cropping systems. This review focus on the specific drought tolerance characteristics in grass species and application of prevalent classical and molecular methods for genetic improvement of them to drought stress. Generally, grass species adapt to drought stress by utilizing more than one strategy including of changes in the root growth, photosynthetic pigments, activation of antioxidant enzymes, and accumulation of compatible osmolytes. They also have other specific characteristics consisted of summer dormancy, drought recovery, and persistence, which lead to drought adaptation after prolonged drought. Studies on different grasses, indicated that most of above mentioned traits usually have positive correlation with drought tolerance. Also, high heritability has been reported for most of them in different grasses. Therefore, an effective index might be considering in identification of drought tolerance genotypes. Recently, high-throughput imaging phenotyping and advanced molecular techniques such as genotyping-by-sequencing (GBS), RNA sequencing, genome-wide association study, and genome editing help conventional breeding methods to increase the accuracy, selection efficiency, genetic gains, and speed of breeding programs for developing drought tolerant cultivars.

Allelopathic Effect of Aqueous Extracts of Grass Genotypes on Eruca Sativa L.

The aim of the current research is to evaluate the allelopathic activity of fifty grass genotypes from different species and to identify phenolic compounds in the genotypes that have the highest allelopathic activity and inhibitory effect on Eruca sativa L. (Rocket). Aqueous extract was prepared from the leaves of grass genotypes in different concentrations and its effect on germination and growth of E. sativa L. was measured. According to the results, the type of genotype and the concentration of the extract significantly decreased the percentage of germination, hypocotyl length, radicle length, and dry weight of E. sativa L. seedlings. Increasing the concentration of the extract resulted in a decrease in germination and growth of seedlings. The genotypes of Festulolium (Festulolium) (GR 5009, GR 1692, GR 5004) had the most inhibitory effect on the growth of E. sativa L. Also, among the genotypes studied, two genotypes (DG-M) and (DG-P) of Dactylis glomerata L. (orchardgrass) species showed the least allelopathic activity. The results of HPLC-MS indicated nine phenolic compounds including caffeic acid, syringic acid, vanillic acid, p-coumaric acid, ferulic acid, apigenin acid, chlorogenic acid, 4-hydroxybenzoic acid, and gallic acid. The phenolic compound most present in the aqueous extract was caffeic acid. However, phenolic compounds derived from Festulolium genotypes showed the greatest allelopathic action on the growth parameters of E. sativa L. The aqueous extracts of the Festulolium genotypes can be considered valid systems of sustainable weed control thanks to the phytocomplex rich in phenols.

Drought stress memory in orchard grass and the role of marker-based parental selection for physiological and antioxidant responses.

Drought stress memory occurring in some plants plays a crucial role in their adaptation to unfavorable conditions. However, in open-pollinated plants, this phenomenon is assumed to be affected by population plasticity resulting from kind and level of diversity and inbreeding depression. Physiological perspectives of drought stress memory in four synthetic poly-crossed populations (groups) of orchard grass (Dactylis glomerata) constructed from parental genotypes with contrasting levels (narrow and wide) of molecular and morphological genetic variation were assessed. Populations of two generations (Syn1 and Syn2) were developed and were subjected to three moisture treatments, including normal irrigation (C), primary mild stress-secondary intense stress (D1D2), and secondary intense stress (D2). Pre-exposure to drought significantly improved the mean values of leaf water, chlorophyll, proline, and ascorbate peroxidase compared to intense stress, leading to more effective memory responses. Superiority of groups with high levels of molecular diversity for most traits, suggesting that the molecular genetic distance among parents is an effective predictor of progeny performance. The results indicated that the fitness of progenies of the four polycross groups declines significantly from Syn1 to Syn2, however the magnitude of observed inbreeding depends on the level of diversity and moisture conditions. We propose a hypothesis that underscores the interplay between genetic diversity among parents and drought stress memory providing valuable insights for developing new synthetic varieties in open-pollinated grasses. Specifically, we posit that higher molecular diversity among parental genotypes enhances the potential for robust drought stress memory, thereby contributing to improved progeny fitness under unfavorable conditions.

Drought stress memory in a germplasm of synthetic and common wheat: antioxidant system, physiological and morphological consequences.

Plants have evolved mechanisms of adaptation to fluctuations in their environmental conditions that have been given the term "stress memory". Synthetic wheat offers new hope for breeders to restore useful genes lost during the genetic bottleneck. We aimed to test whether drought priming and seed priming could improve drought tolerance in a diverse germplasm of synthetic and common wheat under field conditions. In this research, 27 wheat genotypes (including 20 synthetics, 4 common local and 3 common exotic bread wheat) were field evaluated under four water environments. These treatments included: 1) normal condition (N), plants were irrigated when 40% of the total available soil water was depleted from the root-zone, 2) seed priming-secondary stress (SD2), only water stress was applied at anthesis when 90% of the total available soil water was depleted and seeds were planted for evaluating, 3) primary stress- secondary stress (D1D2), primary water stress was applied at jointing stage when 70% of the total available soil water was depleted then secondary water stress was applied at the anthesis stage when 90% of the total available soil water was depleted, and 4) secondary stress (D2) only water stress was applied at the anthesis when 90% of the total available soil water was depleted. Our results indicated that improved efficient enzymatic antioxidant system leads to less yield reduction in D1D2 treatment. However, the positive effects of drought priming were more pronounced in drought primed (D1D2) than seed primed treatment (SD2). Synthetic wheat genotypes had a significant superiority in terms of yield, yield components and drought tolerance compared to common wheat genotypes. Nevertheless, the response of genotypes to stress memory was very different. Drought sensitive genotypes had better response to stress memory. Superior genotypes were identified as high yield and drought tolerant genotypes which can be used for future studies.

Plant functional trait responses to cope with drought in seven cool-season grasses.

In semi-arid and arid regions, the selection of suitable grass species with high-yield production, tolerance to drought stress, and potential for recovery from drought is of special importance. Despite extensive research in cool-season grasses, inter-species differences in post-drought recovery, persistence, survival, and summer dormancy and their relationship with drought tolerance need more investigation. In the present study, 28 diverse genotypes belonged to seven cool-season grass species, including Festuca arundinacea (tall fescue), Festuca pratensis (meadow fescue), Festuca ovina (sheep fescue), Festuca rubra (red fescue), Lolium perenne (perennial ryegrass), Lolium multiflorum (Italian ryegrass) and Lolium × hybridum were evaluated during 2016-2019 under three irrigation regimes (normal, mild, and intense drought stress). Then in the fourth year (on August 2019), irrigation was withheld at all previous irrigation regimes for two months during summer, and then species were re-irrigated to study the effect of prolonged drought conditions. A wide range of genetic diversity was detected in all the measured traits among and within species in response to different irrigation levels. Recurrent drought stress decreased forage productivity, post-drought recovery, and survival in all grass species. Among the studied species, tall fescue had higher forage production, drought tolerance, survival, recovery rate, and persistence. Sheep fescue had low forage production and recovery after drought. Drought tolerance (based on stress tolerance score, STS) was highly associated with forage yield and post-drought recovery and partially with summer dormancy under both mild and intense drought stress conditions. This indicated that selection based on higher STS would lead to choosing genotypes with better recovery after prolonged drought. Superior species and preferable genotypes for forage use from species Festuca arundinacea and for turf application from species Festuca arundinacea, Lolium perenne and Lolium × hybridum were identified across different water environments for future programs.

Characterization of fennel germplasm for physiological persistence and drought recovery: Association with biochemical properties.

In the perennial medicinal plant fennel (Foeniculum vulgare), persistence over years, production stability, and successful post-drought recovery are as important as plant productivity. Characterization of productivity, post-drought recovery, persistency and their association with phytochemical properties has not yet been performed in fennel. In this study, 64 fennel genotypes (from 23 different countries) from four subspecies/varieties including F. vulgare ssp. piperitum, F. vulgare var. vulgare and F. vulgare var. dulce and F. vulgare var. azoricum were evaluated in the field over four years (2015-2018), then was assessed for post-drought recovery over the next two years (2019-2020). High genotypic variation was observed among genotypes and subspecies. Based on the GC-MS analysis, trans-anethole (22.4-90.6%), estragole (2.1-25.8%), fenchone (4.9-19.8%), and limonene (0.5-11.9%) are major components in the essential oils of the studied germplasm. The highest persistence and the lowest average post-drought recovery belonged to ssp. piperitum. This subspecies also had the highest amount of limonene, fenchone, and estragole and the lowest amount of trans-anethole compared to other subspecies. The highest essential oil content and stability was observed in var. dulce. Seed yield and persistence were found to be negatively associated, suggesting that selection for more productivity may indirectly promote less persistent genotypes. Post-drought recovery was negatively associated with two main phytochemical compounds of essential oil, fenchone and estragole, but positively associated with trans-anethole. Persistence had negative correlation with estragole content. Results indicated that indirect selection for post-drought recovery and persistence may be possible through selection for phytochemical properties. The negative correlation between some essential oil components may indicate that some are isomers, which can limit the ability to select for certain combinations. Superior genotypes identified in this study can be used to construct populations for future studies and as parents of crosses to develop new varieties.

Association analysis revealed loci linked to post-drought recovery and traits related to persistence of smooth bromegrass (Bromus inermis).

Association analysis has been proven as a powerful tool for the genetic dissection of complex traits. This study was conducted to identify association of recovery, persistence, and summer dormancy with sequence related amplified polymorphism (SRAP) markers in 36 smooth bromegrass genotypes under two moisture conditions and find stable associations. In this study, a diverse panel of polycross-derived progenies of smooth bromegrass was phenotyped under normal and water deficit regimes for three consecutive years. Under water deficit, dry matter yield of cut 1 was approximately reduced by 36, 39, and 37% during 2013, 2014, and 2015, respectively, compared with the normal regime. For dry matter yield of cut 2, these reductions were approximately 38, 60, and 56% in the same three consecutive years relative to normal regime. Moreover, water deficit decreased the RY and PER of the genotypes by 35 and 28%, respectively. Thirty primer combinations were screened by polymerase chain reaction (PCR). From these, 541 polymorphic bands were developed and subjected to association analysis using the mixed linear model (MLM). Population structure analysis identified five main subpopulations possessing significant genetic differences. Association analysis identified 69 and 46 marker-trait associations under normal and water deficit regimes, respectively. Some of these markers were associated with more than one trait; which can be attributed to pleiotropic effects or tightly linked genes affecting several traits. In normal and water-deficit regimes, these markers could potentially be incorporated into marker-assisted selection and targeted trait introgression for the improvement of drought tolerance of smooth bromegrass.

Potentials of synthetic hexaploid wheats to improve drought tolerance.

Synthetic hexaploid wheat-derived lines (SHW-DL) offers new hope for breeders to restore genes lost during the evolutionary bottleneck. The study of adaptability, variation, and the possibility of selection in SHW-DL for drought tolerance is poorly understood in arid environments. The potential of 184 SHW-DL and their variation for agro-morphological traits were assessed under normal and water stress conditions for 2 years. The mean values of grain yield (YLD) varied from 683.9 g/m2 (water stress) to 992.1 g/m2 (normal conditions). Grain yield decreased by 64 and 71% under water stress in the two growing seasons. High genotypic variation was found for measured traits and drought tolerance. Heritability ranged from 19 (harvest index) to 47% (spike length), whereas grain yield indicated a moderate heritability (32%). Using the assessment of the interrelationship of traits, hectoliter (a quality trait) was correlated with drought tolerance and stability indices. Therefore, it can be considered as an important trait to select drought tolerant genotypes. In the following, the priority of yield components entering the regression model was different in two moisture conditions suggesting different strategies in indirect selection programs to improve yield. Spike m-2 and grain spike-1 indirectly and negatively affected yield through thousand-grain weight (TGW) under normal and water stress conditions, respectively. Furthermore, SHW-DL compared to ordinary wheat were significantly superior in terms of early maturity, dwarfing, yield, TGW, stem diameter, and harvest index. Overall, our findings suggest that SHW-DL are a valuable source for improving wheat yield and drought tolerance, and indirect selection might be possible to improve these complex traits.

Genetic analysis and physiological relationships of drought response in fennel: Interaction with mating system.

The consequences of water deficit and its interaction with pollination system (deliberate selfing compared with open-pollination) on physiological, agronomic and phytochemical traits are not understood in fennel (Foeniculum vulgare Mill.). A research was started by creating selfed (S1) and half-sib (HS) families on a fennel germplasm in 2018. Populations were studied in the field, applying a normal and a water deficit condition during two years (2019-2020). Considerable genotypic variation was observed within S1 and HS families for all of the evaluated traits, demonstrating that selection for these traits would be successful. Consequences of water deficit were manifested as declined most of the traits; and significantly increased essential oil content, harvest index, and proline content, in both populations. Mandatory selfing reduced the performance of genotypes for most of the traits confirming the existence of inbreeding depression (ID) with higher values for plant dry weight, seed yield, essential oil content, and number of umbelets per umbel. In S1 population, some of the studied traits had higher heritability estimates under normal condition and some of them showed higher heritability under water deficit. Positive relationship between GCA and STI in OP population indicated that it is possible to identify genotypes having high values of combining ability and drought tolerance. Results of the present study suggest that physiological traits cannot be used as an indicator to distinguish drought-tolerant genotypes in S1 progenies, whereas in OP progenies Chl a, Chl b, TChl, CAR, PRO, and RWC, which had significant correlations with drought tolerance, may be used for this purpose. Based on the results contrasting genotypes were identified, which can be used to develop mapping populations for genome studies of drought tolerance and physiological traits of this species in future studies.

Genetic parameters and selection in full-sib families of tall fescue using best linear unbiased prediction (BLUP) analysis.

BACKGROUND: Better understanding of genetic structure of economic traits is crucial for identification and selection of superior genotypes in specific breeding programs. Best linear unbiased prediction (BLUP) is the most efficient method in this regard, which is poorly used in forage plant breeding. The present study aimed to assess genetic variation, estimate genetic parameters, and predict breeding values of five essential traits in full sib families (recognized by EST-SSR markers) of tall fescue using REML/BLUP procedure. METHOD: Forty-two full-sib families of tall fescue (included of 120 individual genotypes), recognized by EST-SSR markers along with twenty-one their corresponding parental genotypes were assessed for biomass production and agro-morphological traits at three harvests (spring, summer, and autumn) in the field during 4 years (2017-2020). RESULTS: Considerable genotypic variability was observed for all traits. Low narrow-sense heritability (h2n) for dry forage yield (DFY) at three harvest indicates that non-additive gene actions may play an important role in the inheritance of this trait. Higher h2n of yield related traits and flowering time and also significant genetic correlation of these traits with forage yield, suggests that selection based on these traits may lead to indirect genetic improvement of DFY. CONCLUSION: Our results showed the adequacy of REML/BLUP procedure for identification and selection of preferable parental genotypes and progenies with higher breeding values for future breeding programs such as variety development in tall fescue. Parental genotypes 21 M, 1 M, and 20 L were identified as superior and stable genotypes and could also produce the best hybrid combinations when they were mostly used as maternal parent.

Changes in activities of antioxidant enzymes in radish (Raphanus sativus) seedlings in response to allelopathic effect of safflower (Carthamus tinctorius)

Abstract ROS (Reactive Oxygen Species) production is a usual plant reaction to environmental stresses such as allelopathy. Plants possess antioxidant enzymes to scavenge cells and resist against the ROS. This study was conducted to evaluate changes in antioxidant enzymes (CAT, GPX, APX) in radish seedlings in response to allelopathic effect of safflower root and shoot residues grown under normal irrigation and drought stress. Safflower allelopathic effect led to an increase in antioxidant enzymes activities. GPX activity increased more than CAT and APX. Radish seedlings exposed to safflower residue grown under drought stress showed more antioxidant enzymes activities. Root residues enhanced the activities of antioxidant enzymes greater than shoot. Seedlings exposed to root residues grown under drought stress had the highest level of antioxidant enzymes activities.

Evidence for the Accumulation of Nonsynonymous Mutations and Favorable Pleiotropic Alleles During Wheat Breeding.

Plant breeding leads to the genetic improvement of target traits by selecting a small number of genotypes from among typically large numbers of candidate genotypes after careful evaluation. In this study, we first investigated how mutations at conserved nucleotide sites normally viewed as deleterious, such as nonsynonymous sites, accumulated in a wheat, Triticum aestivum, breeding lineage. By comparing a 150 year old ancestral and modern cultivar, we found recent nucleotide polymorphisms altered amino acids and occurred within conserved genes at frequencies expected in the absence of purifying selection. Mutations that are deleterious in other contexts likely had very small or no effects on target traits within the breeding lineage. Second, we investigated if breeders selected alleles with favorable effects on some traits and unfavorable effects on others and used different alleles to compensate for the latter. An analysis of a segregating population derived from the ancestral and modern parents provided one example of this phenomenon. The recent cultivar contains the Rht-B1b green revolution semi-dwarfing allele and compensatory alleles that reduce its negative effects. However, improvements in traits other than plant height were due to pleiotropic loci with favorable effects on traits and to favorable loci with no detectable pleiotropic effects. Wheat breeding appears to tolerate mutations at conserved nucleotide sites and to only select for alleles with both favorable and unfavorable effects on traits in exceptional situations.

Expression pattern of salt tolerance-related genes in Aegilops cylindrica.

Aegilops cylindrica, a salt-tolerant gene pool of wheat, is a useful plant model for understanding mechanism of salt tolerance. A salt-tolerant USL26 and a salt-sensitive K44 genotypes of A. cylindrica, originating from Uremia Salt Lake shores in Northwest Iran and a non-saline Kurdestan province in West Iran, respectively, were identified based on screening evaluation and used for this work. The objective of the current study was to investigate the expression patterns of four genes related to ion homeostasis in this species. Under treatment of 400 mM NaCl, USL26 showed significantly higher root and shoot dry matter levels and K+ concentrations, together with lower Na+ concentrations than K44 genotype. A. cylindrica HKT1;5 (AecHKT1;5), SOS1 (AecSOS1), NHX1 (AecNHX1) and VP1 (AecVP1) were partially sequenced to design each gene specific primer. Quantitative real-time PCR showed a differential expression pattern of these genes between the two genotypes and between the root and shoot tissues. Expressions of AecHKT1;5 and AecSOS1 was greater in the roots than in the shoots of USL26 while AecNHX1 and AecVP1 were equally expressed in both tissues of USL26 and K44. The higher transcripts of AecHKT1;5 in the roots versus the shoots could explain both the lower Na+ in the shoots and the much lower Na+ and higher K+ concentrations in the roots/shoots of USL26 compared to K44. Therefore, the involvement of AecHKT1;5 in shoot-to-root handover of Na+ in possible combination with the exclusion of excessive Na+ from the root in the salt-tolerant genotype are suggested.