Ionic and osmotic relations in quinoa (Chenopodium quinoa Willd.) were studied by exposing plants to six salinity levels (0–500 mM NaCl range) for 70 d. Salt stress was administered either by pre-mixing of the calculated amount of NaCl with the potting mix before seeds were planted or by the gradual increase of NaCl levels in the irrigation water. For both methods, the optimal plant growth and biomass was achieved between 100 mM and 200 mM NaCl, suggesting that quinoa possess a very efficient system to adjust osmotically for abrupt increases in NaCl stress. Up to 95% of osmotic adjustment in old leaves and between 80% and 85% of osmotic adjustment in young leaves was achieved by means of accumulation of inorganic ions (Na+, K+, and Cl–) at these NaCl levels, whilst the contribution of organic osmolytes was very limited. Consistently higher K+ and lower Na+ levels were found in young, as compared with old leaves, for all salinity treatments. The shoot sap K+ progressively increased with increased salinity in old leaves; this is interpreted as evidence for the important role of free K+ in leaf osmotic adjustment under saline conditions. A 5-fold increase in salinity level (from 100 mM to 500 mM) resulted in only a 50% increase in the sap Na+ content, suggesting either a very strict control of xylem Na+ loading or an efficient Na+ removal from leaves. A very strong correlation between NaCl-induced K+ and H+ fluxes was observed in quinoa root, suggesting that a rapid NaCl-induced activation of H+-ATPase is needed to restore otherwise depolarized membrane potential and prevent further K+ leak from the cytosol. Taken together, this work emphasizes the role of inorganic ions for osmotic adjustment in halophytes and calls for more in-depth studies of the mechanisms of vacuolar Na+ sequestration, control of Na+ and K+ xylem loading, and their transport to the shoot.
Wheat breeding for salinity tolerance has traditionally focussed on Na+ exclusion from the shoot, but its association with salinity tolerance remains tenuous. Accordingly, the physiological significance of shoot Na+ exclusion and maintenance of an optimal K+ : Na+ ratio was re-evaluated by studying NaCl-induced responses in 50 genotypes of bread wheat (Triticum aestivum L.) and durum wheat (Triticum turgidum L. ssp. durum) treated with 150 mM NaCl. Overall, Na+ exclusion from the shoot correlated with salinity tolerance in both species and this exclusion was more efficient in bread compared with durum wheat. Interestingly, shoot sap K+ increased significantly in nearly all durum and bread wheat genotypes. Conversely, the total shoot K+ content declined. We argue that this increase in shoot sap K+ is needed to provide efficient osmotic adjustment under saline conditions. Durum wheat was able to completely adjust shoot sap osmolality using K+, Na+ and Cl–; it had intrinsically higher levels of these solutes. In bread wheat, organic osmolytes must contribute ~13% of the total shoot osmolality. In contrast to barley (Hordeum vulgare L.), NaCl-induced K+ efflux from seedling roots did not predict salinity tolerance in wheat, implying that shoot, not root K+ retention is important in this species.
Two components of salinity stress are a reduction in water availability to plants and the formation of reactive oxygen species. In this work, we have used quinoa (Chenopodium quinoa), a dicotyledonous C3 halophyte species displaying optimal growth at approximately 150 mM NaCl, to study mechanisms by which halophytes cope with the afore-mentioned components of salt stress. The relative contribution of organic and inorganic osmolytes in leaves of different physiological ages (e.g. positions on the stem) was quantified and linked with the osmoprotective function of organic osmolytes. We show that the extent of the oxidative stress (UV-B irradiation) damage to photosynthetic machinery in young leaves is much less when compared with old leaves, and attribute this difference to the difference in the size of the organic osmolyte pool (1.5-fold difference under control conditions; sixfold difference in plants grown at 400 mM NaCl). Consistent with this, salt-grown plants showed higher Fv/Fm values compared with control plants after UV-B exposure. Exogenous application of physiologically relevant concentrations of glycine betaine substantially mitigated oxidative stress damage to PSII, in a dose-dependent manner. We also show that salt-grown plants showed a significant (approximately 30%) reduction in stomatal density observed in all leaves. It is concluded that accumulation of organic osmolytes plays a dual role providing, in addition to osmotic adjustment, protection of photosynthetic machinery against oxidative stress in developing leaves. It is also suggested that salinity-induced reduction in stomatal density represents a fundamental mechanism by which plants optimize water use efficiency under saline conditions.
Improved soybean cultivars have been adapted to grow at a wide range of latitudes, enabling expansion of cultivation worldwide. However, the genetic basis of this broad adaptation is still not clear. Here, we report the identification of GmPRR3b as a major flowering time regulatory gene that has been selected during domestication and genetic improvement for geographic expansion. Through a genome-wide association study of a diverse soybean landrace panel consisting of 279 accessions, we identified 16 candidate quantitative loci associated with flowering time and maturity time. The strongest signal resides in the known flowering gene E2, verifying the effectiveness of our approach. We detected strong signals associated with both flowering and maturity time in a genomic region containing GmPRR3b. Haplotype analysis revealed that GmPRR3b H6 is the major form of GmPRR3b that has been utilized during recent breeding of modern cultivars. mRNA profiling analysis showed that GmPRR3b H6 displays rhythmic and photoperioddependent expression and is preferentially induced under long-day conditions. Overexpression of GmPRR3b H6 increased main stem node number and yield, while knockout of GmPRR3b H6 using CRISPR/ Cas9 technology delayed growth and the floral transition. GmPRR3b H6 appears to act as a transcriptional repressor of multiple predicted circadian clock genes, including GmCCA1a, which directly upregulates J/GmELF3a to modulate flowering time. The causal SNP (Chr12:5520945) likely endows GmPRR3b H6 a moderate but appropriate level of activity, leading to early flowering and vigorous growth traits preferentially selected during broad adaptation of landraces and improvement of cultivars.
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