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Osmoregulation and salinity effects on the expression and activity of Na+,K(+)-ATPase in the gills of European sea bass, Dicentrarchus labrax (L.).

The European sea bass, Dicentrarchus labrax, tolerates salinities ranging from freshwater (FW) to hypersaline conditions. In two experiments, we analysed changes in plasma ions, muscle water content (MWC), gill Na+,K(+)-ATPase activity, and alpha-subunit mRNA expression during the course of acclimation from 15 ppt salt water to FW or high salinity seawater (HSSW). In Experiment 1, fish (6.2 +/- 1.1 g) were acclimated from 15 ppt to either FW, 5, 15, 25, 50, or 60 ppt SW and sampled after 10 days. Gill Na+,K(+)-ATPase activity was stimulated in FW- and in 50 and 60 ppt SW-groups relative to the 15 ppt control group. In Experiment 2, subgroups of fish (89 +/- 7 g) were transferred from 15 ppt SW to FW or 50 ppt SW, and sampled 1, 2, 4, and 10 days later. Plasma osmolality, [Na+] and [Cl-] decreased in the FW-group and increased in the HSSW-group one day after transfer and lasting until day 10. This was accompanied by a pronounced increase in MWC in the FW-group and an insignificant decrease in the HSSW-group. The plasma [Na+]:[Cl-]-ratio increased markedly in the FW-group and decreased slightly in the HSSW-group, suggesting acid-base balance disturbances after transfer. Gill Na+,K(+)-ATPase activity was unchanged in 15 ppt SW but doubled in FW- and HSSW-groups after transfer. In both groups, this was preceded by a 2- to 5-fold elevation of the gill alpha-subunit Na+,K(+)-ATPase mRNA level. Thus increased expression of alpha-subunit mRNA is part of the molecular mechanism of both FW and SW acclimation in sea bass. Gill Na+,K(+)-ATPase Na(+)-, K(+)-, and ouabain-affinity were similar in fish acclimated to FW, 15 ppt, and HSSW, suggesting that identical isoforms of the catalytic subunit of the enzyme are expressed irrespective of salinity.

Adaptation, Physiological↗

Tandem gene duplication facilitates intertidal adaptation in atypical mangrove plants.

Mangrove plants, originating from inland ancestors, have independently adapted to extreme intertidal zones characterized by salt and hypoxia stress. While typical mangroves exhibit specialized phenotypes, like viviparous seeds and salt secretion, atypical clades that have thrived without such traits are particularly suitable for exploring the molecular and physiological basis underlying plant adaptation to intertidal zones. We assembled a chromosome-level genome of an atypical mangrove, Scyphiphora hydrophylacea, the only mangrove species in Gentianales. Similar to other mangroves, S. hydrophylacea colonized intertidal zones during climatic optimum periods of sea-level rise. Despite lacking recent whole-genome duplications (WGDs), its genome acquired extensive tandem gene duplications (TDs), leading to the rapid expansion of key salt- and hypoxia-related genes. Transcriptome data further corroborated that TD-driven gene expansions contribute to stress tolerance. Specifically, the expansion of genes involved in cation transmembrane transport, osmotic regulation, and oxidative stress response may enhance salinity tolerance, and the expansion of signal transduction and energy metabolism genes in hypoxia-response pathways may confer waterlogging tolerance. Therefore, in the absence of large-scale gene duplication, the rapid expansion of core genes involved in salt and hypoxia tolerance through tandem duplication may represent a key force driving the adaptation of atypical mangroves. These findings also provide valuable insights for crop improvement strategies aimed at enhancing environmental resilience while maintaining phenotypic stability.

Gene Duplication↗

pH tolerances and regulatory abilities of freshwater and euryhaline Aedine mosquito larvae.

The pH regulatory abilities of two members of the mosquito tribe Aedini, known to have dramatically different saline tolerances, are investigated. The freshwater mosquito Aedes aegypti and the euryhaline Ochlerotatus taeniorhynchus tolerate very similar pH ranges. Both species complete larval development in waters ranging from pH 4 to pH 11, but naïve larvae always die in water of pH 3 or 12. Across the pH range 4-11, the hemolymph pH of O. taeniorhynchus is maintained constant while that of A. aegypti varies by 0.1 pH units. The salt composition of the water (3.5 g l(-1) sea salt, 3.5 g l(-1) NaCl, or nominally salt-free) has no effect on the range of pH tolerated by A. aegypti. In both species, the effects of pH on larval growth and development are minor in comparison with the influence of species and sex. Acclimation of A. aegypti to pH 4 or 11 increases survival times in pH 3 or 12, respectively, and allows a small percentage of larvae to pupate successfully at these extreme pH values. Such acclimation does not compromise survival at the other pH extreme.

Acclimatization↗

Morphological and Physiological Responses of Hawaiian Hibiscus tiliaceus Populations to Light and Salinity.

Hibiscus tiliaceus (Hau) is a pantropical mangrove associate that usually occurs in coastal ecosystems where substrate salinity is relatively high, but it also inhabits upland habitats in Hawaii. Cuttings from three populations on the island of Oahu, Hawaii, were collected and grown in the glasshouse under two levels of substrate salinity (0 and 335 mOsm kg-1) and three light treatments (0%, 50%, and 90% shade). Photosynthetic gas exchange, biomass allocation, and accumulation were studied in relation to salinity and light. Salinity reduced net CO2 assimilation in the upland population but had no effect or stimulated photosynthesis in the coastal populations, whereas increasing salinity decreased stomatal conductance in all populations and therefore increased water-use efficiency. The degree to which photosynthesis was inhibited by salinity was inversely proportional to the salinity of the source population, indicating a loss of salinity tolerance in upland plants. Light had a stronger effect on leaf area ratio (LAR) and leaf mass per area (LMA), whereas salinity had a stronger effect on leaf water content, internode length, and plant biomass. Salinity reduced total new biomass by 58%, 50%, and 34% in full sun, 50% shade, and 90% shade, respectively, but this response did not differ between populations. Salinity reduced the photosynthesis, but not growth, of upland plants because increased allocation to photosynthetic tissue increased LAR to compensate for inhibition of photosynthesis by salinity.

Journal Article↗

Investigations on cyanobacterial diversity in a shallow estuary (Southern Baltic Sea) including genes relevant to salinity resistance and iron starvation acclimation.

The cyanobacterial diversity in the pelagic of a shallow estuary at the Southern Baltic Sea has been investigated by a combination of classical morphological data and a polymerase chain reaction (PCR)-based molecular approach. The aim of the study was to investigate possible changes in the composition of the cyanobacterial community along the salinity and nutrient gradients. For this purpose partial gene sequences of cyanobacterial 16S rDNA and of two functional genes (ggpS- salinity tolerance marker, isiA- iron starvation marker) were amplified and compared with total community DNA. Random distribution of ggpS genotypes along the salinity gradient suggests that synthesis of the osmolyte glucosylglycerol is not restricted to higher salinity sampling sites. Most of the isiA sequences formed a new homogenous cluster in a phylogenetic tree, which indicates that the indigenous cyanobacterial community comprises a group of unknown species. Minimum iron concentrations, which can activate isiA transcription in model cyanobacteria, occurred at a few sampling sites with high phytoplankton biomass and moderate salinity. Nevertheless, isiA expression could be detected at all sampling sites, which indicated restricted iron supply to cyanobacterial phytoplankton in summer.

Acclimatization↗

A deep metagenomic atlas of Qinghai-Xizang Plateau lakes reveals their microbial diversity and salinity adaptation mechanisms.

The Qinghai-Xizang Plateau (QXP), harboring the planet's highest density of plateau lakes, offers an exceptional biogeographic environment for studying extremophilic microbial communities and their adaptation to salinity. Through deep metagenomic sequencing, we construct the Qinghai-Xizang Lake Sediment Genome (QXLSG) catalog, a high-resolution genomic catalog comprising 5,866 metagenome-assembled genomes (MAGs), 58.16 million non-redundant protein encoding genes, and 19,008 biosynthetic gene clusters. Notably, 80.78% of the 2,742 species-level MAGs represent undescribed taxa, significantly expanding the known microbial diversity. Salinity emerges as the primary environmental factor influencing microbial community. Functional annotation highlights that the "salt-out" strategy, particularly the uptake of glycine betaine, is the main mechanism for salinity tolerance. This strategy is prevalent in both hypersaline lake communities and the dominant microbial phyla. Overall, this study provides a crucial genetic resource for future bioprospecting and deepens our understanding of the fundamental mechanisms of microbial adaptation to extreme saline environments.

Lakes↗

Effect of water quality on yield of sugar beet and sweet sorghum.

To study the effects of quality of water on soil and plant, an experiment was conducted at Rudashat Drainage and Reclamation Experiment Station in 1999. Four irrigation water salinities (2, 5, 8 and 11 ds m(-1)) and two sugar crops (sugar beet and sweet sorghum) were used in this experiment. The results showed that under the same water quality, sweet sorghum used 2700 cubic meter per hectare less water than sugar beet. As the quality of irrigation water decreased, the soil salinity and exchangeable sodium percent increased which caused yield reduction for both plants. Sugar beet by accumulating Na and Cl in its leaves tolerated salinity but its usage as a forage crop caused some limitations, whereas sweet sorghum by not accumulating Na and Cl escape salinity and it can be used as a forage crop without any limitation.

Agriculture↗

Male and female gametophyte selection of barley for salt tolerance.

The contribution of the gametophyte in the inheritance of salt tolerance was studied by crossing F3 and BC1 hybrids of the tolerant cultivars Rannii 1 and Pirkka with the sensitive cultivar K-30356. The third generation shows that the progenies of heterozygous plants grown in saline conditions show elevated salinity tolerance. Again, a comparison of the BC1 hybrid progenies shows that the male and female gametophytes contribute to the inheritance of salt tolerance. Gametic selection is maximally efficient during the formation of the female gametophyte and the germination of pollen grains on the stigma.

Alleles↗

Response of biotic communities to salinity changes in a Mediterranean hypersaline stream.

BACKGROUND: This study investigates the relationship between salinity and biotic communities (primary producers and macroinvertebrates) in Rambla Salada, a Mediterranean hypersaline stream in SE Spain. Since the 1980's, the mean salinity of the stream has fallen from about 100 g L(-1) to 35.5 g L(-1), due to intensive irrigated agriculture in the watershed. Furthermore, large dilutions occur occasionally when the water irrigation channel suffers cracks. RESULTS: Along the salinity gradient studied (3.5-76.4 g L(-1)) Cladophora glomerata and Ruppia maritima biomass decreased with increasing salinity, while the biomass of epipelic algae increased. Diptera and Coleoptera species dominated the community both in disturbed as in re-established conditions. Most macroinvertebrates species found in Rambla Salada stream are euryhaline species with a broad range of salinity tolerance. Eight of them were recorded in natural hypersaline conditions (approximately 100 g L(-1)) prior to important change in land use of the watershed: Ephydra flavipes, Stratyomis longicornis, Nebrioporus ceresyi, N. baeticus, Berosus hispanicus, Enochrus falcarius, Ochthebius cuprescens and Sigara selecta. However, other species recorded in the past, such as Ochthebius glaber, O. notabilis and Enochrus politus, were restricted to a hypersaline source or absent from Rambla Salada. The dilution of salinity to 3.5-6.8 g L(-1) allowed the colonization of species with low salininty tolerance, such as Melanopsis praemorsa, Anax sp., Simulidae, Ceratopogonidae and Tanypodinae. The abundance of Ephydra flavipes and Ochthebius corrugatus showed a positive significant response to salinity, while Anax sp., Simulidae, S. selecta, N. ceresyi, N. baeticus, and B. hispanicus showed significant negative correlations. The number of total macroinvertebrate taxa, Diptera and Coleoptera species, number of families, Margalef's index and Shannon's diversity index decreased with increasing salinity. However, the rest of community parameters, such as the abundance of individuals, evenness and Simpson's index, showed no significant response to changes in salinity. Classification and ordination analysis revealed major differences in macroinvertebrate community structure between hypersaline conditions (76.4 g L(-1)) and the rest of the communities observed at the lower salinity levels, and revealed that below approximately 75 g L(-1), dissimilarities in the communities were greater between the two habitats studied (runs and pools) than between salinity levels. CONCLUSION: Salinity was the first factor determining community composition and structure in Rambla Salada stream followed by the type of habitat.

Journal Article↗

Effect of chronic administration of dihydropyridine Ca2+ channel ligands on sufentanil-induced tolerance to mu- and kappa-opioid agonists in the guinea pig ileum myenteric plexus.

The present investigation was aimed at elucidating if the entry of Ca2+ plays a role in the development of tolerance to mu- and kappa-opioid agonists in the guinea pig ileum myenteric plexus. For this purpose, the influence of the L-type Ca2+ channel modulators nimodipine (Ca2+ blocker) and Bay K 8644 (Ca2+ activator) on the expression of tolerance to the inhibitory effects of mu- and kappa-opioid agonists in the ileum of guinea pigs rendered tolerant to sufentanil was investigated. Chronic perfusion of guinea pigs with nimodipine (2 micrograms/microliter/h for 7 days) or Bay K 8644 (0.5 microgram/microliter/h for 7 days) did not cause any modification of the height of contractions induced by electrical stimulation of the myenteric plexus-longitudinal muscle (MPLM) strip from naive animals. Tolerance to sufentanil (a selective mu-agonist) was induced by s.c. implantation of osmotic minipumps for 7 days, which deliver at 2 micrograms/microliter/h. Control groups received saline. Tolerance to sufentanil as well as to U-50,488H (selective kappa-agonist) was observed following chronic treatment with sufentanil and was revealed as a rightward shift of the concentration-response curves. Chronic perfusion of guinea pigs with the Ca2+ antagonist nimodipine concurrently with chronic sufentanil, markedly blocked the expression of tolerance to sufentanil, as well as the cross-tolerance between sufentanil and U-50,488H. On the contrary, when guinea pigs were perfused with the Ca2+ agonist Bay K 8644 concurrently with sufentanil, it enhanced the magnitude of tolerance to both sufentanil and U-50,488H. These results suggest that, in guinea pig ileum, chronic exposure to opioids may involve the activation of L-type Ca2+ channel, which would indicate that intracellular Ca2+ may be one of the final pathways through which myenteric neurons adapt to the chronic opioid exposure.

3,4-Dichloro-N-methyl-N-(2-(1-pyrrolidinyl)-cycloh↗

The cytosolic Na+ : K+ ratio does not explain salinity-induced growth impairment in barley: a dual-tracer study using 42K+ and 24Na+.

It has long been believed that maintenance of low Na+ : K+ ratios in the cytosol of plant cells is critical to the plant's ability to tolerate salinity stress. Direct measurements of such ratios, however, have been few. Here we apply the non-invasive technique of compartmental analysis, using the short-lived radiotracers 42K+ and 22Na+, in intact seedlings of barley (Hordeum vulgare L.), to evaluate unidirectional plasma membrane fluxes and cytosolic concentrations of K+ and Na+ in root tissues, under eight nutritional conditions varying in levels of salinity and K+ supply. We show that Na+ : K+ ratios in the cytosol of root cells adjust significantly across the conditions tested, and that these ratios are poor predictors of the plant's growth response to salinity. Our study further demonstrates that Na+ is subject to rapid and futile cycling at the plasma membrane at all levels of Na+ supply, independently of external K+, while K+ influx is reduced by Na+, from a similar baseline, and to a similar extent, at both low and high K+ supply. We compare our results to those of other groups, and conclude that the maintenance of the cytosolic Na+ : K+ ratio is not central to plant survival under NaCl stress. We offer alternative explanations for sodium sensitivity in relation to the primary acquisition mechanisms of Na+ and K+.

Cytosol↗

The effect of temperature on juvenile Mozambique tilapia hybrids (Oreochromis mossambicus x O. urolepis hornorum) exposed to full-strength and hypersaline seawater.

The effects of temperature on the salinity tolerance of Mozambique-Wami tilapia hybrids (Oreochromis mossambicus x O. urolepis hornorum) were investigated by transferring 35 g/l, 25 degrees C-acclimated fish to 35, 43, 51 or 60 g/l salinity at 15, 25 or 35 degrees C for 24 h, and by assaying gill tissue for branchial Na(+), K(+)-ATPase activity at the three temperatures after acclimating the fish to 15, 25 or 35 degrees C for 2 weeks. Tilapia survived all salinities at 25 and 35 degrees C; however, at 15 degrees C, mortality was 85.7% and 100% in the 51 g/l and 60 g/l groups, respectively. There was a significant interaction between temperature and salinity, as plasma osmolality, [Na(+)] and [Cl(-)] were significantly increased at 51 and 60 g/l salinity in 35 degrees C water (P<0.001). Additionally, muscle water content was significantly reduced at 43 g/l, 15 degrees C relative to pre-transfer values (P<0.001). Branchial Na(+), K(+)-ATPase activity was reduced at 15 degrees C regardless of acclimation temperature, and 25 degrees C-acclimated gill tissue did not show an increase in activity when assayed at 35 degrees C. Results indicate that the effects of a combined temperature-salinity transfer on plasma osmolality and ion concentrations, as well as muscle water content, are greater than when either challenge is given alone. Additionally, branchial Na(+), K(+)-ATPase activity is altered when assayed at varying temperatures; in the case of 15 degrees C, regardless of acclimation temperature. Our enzyme activity data may indicate the presence of a high temperature isoform of branchial Na(+), K(+)-ATPase enzyme.

Acclimatization↗

Evaluation of anorectal function in patients with tethered cord syndrome: saline enema test and fecoflowmetry.

OBJECT: Disturbance in anorectal function is a major factor restricting the activities of daily living in patients with spinal cord disorders. To detect changes in anorectal motilities due to a tethered spinal cord, anorectal functions were evaluated using a saline enema test and fecoflowmetry before and after patients underwent untethering surgery. METHODS: The bowel functions in five patients with a tethered cord syndrome (TCS) were evaluated by performing a saline enema test and fecoflowmetry. The contractile activity of the rectum, the volume of infused saline tolerated in the rectum, anal canal pressure, and the ability to evacuate rectal content were examined. The characteristic findings in anorectal motility studies conducted in patients with TCS were a hyperactive rectum, diminished rectal saline-retention ability, and diminished maximal flow in saline evacuation. A hyperactive rectum was considered to be a major contributing factor to fecal incontinence. In one asymptomatic patient diminished anal squeezing pressure was exhibited and was incontinent to liquid preoperatively, but recovered after surgery. Two patients who underwent surgery for myeloschisis as infants complained of progressive fecal incontinence when they became adolescents. In one patient fecal incontinence improved but in another patient no improvement was observed after untethering surgery. CONCLUSIONS: Fecodynamic studies allow the detection of neurogenic disturbances of the anorectum in symptomatic and also in asymptomatic patients with TCS. More attention should be paid to the anorectal functions of patients with TCS.

Anal Canal↗

Engineering salt-tolerant Brassica plants: characterization of yield and seed oil quality in transgenic plants with increased vacuolar sodium accumulation.

Transgenic Brassica napus plants overexpressing AtNHX1, a vacuolar Na(+)/H(+) antiport from Arabidopsis thaliana, were able to grow, flower, and produce seeds in the presence of 200 mM sodium chloride. Although the transgenic plants grown in high salinity accumulated sodium up to 6% of their dry weight, growth of the these plants was only marginally affected by the high salt concentration. Moreover, seed yields and the seed oil quality were not affected by the high salinity of the soil. Our results demonstrate the potential use of these transgenic plants for agricultural use in saline soils. Our findings, showing that the modification of a single trait significantly improved the salinity tolerance of this crop plant, suggest that with a combination of breeding and transgenic plants it could be possible to produce salt-tolerant crops with far fewer target traits than had been anticipated.

Journal Article↗

Salinity adaptation in the snake eel Pisoodonophis boro (Ham-buch).

The snake eel Pisoodonophis Boro (Family Ophichthidae) burrows, causing leaks in the bunds and damaging the salterns and paddy fields adjoining estuaries. The animal's capability to withstand brine in salterns and fresh water in paddy fields was determined from laboratory experiments. The eel tolerated salinities ranging from deionized water to 75%. Adaptation to fresh water as well as full strength seawater was completed within 24 h. The concentration of sodium and chloride in the blood varied by 10--20% on transfer from seawater to fresh water. P. boro can survive in paddy fields but not in salterns. However, their occurrence in both is described as a rare phenomenon and not obligatory.

Adaptation, Physiological↗

Differential mRNA transcription during salinity stress in barley.

The molecular and genetic bases of salinity tolerance in plants are not understood. Gene expression at the mRNA level was investigated in a salt-tolerant and a salt-sensitive genotype of barley. Seedlings were exposed to NaCl stress and translatable mRNAs were isolated from root and shoot tissues. A reticulocyte cell-free system was programed with barley mRNAs and the in vitro products were resolved on two-dimensional polyacrylamide gels following isoelectric focusing or nonequilibrium pH gradient gel electrophoresis in the first dimension. The functional mRNAs in unstressed seedlings were qualitatively almost indistinguishable in the two genotypes. However, salinity stress triggered differential transcription of specific mRNAs depending upon genotype and tissue. In roots, 12 new mRNAs were induced that encoded proteins of 21-34 kDa, with a pI range of 6.1-7.7. In shoots, the 9 new mRNAs coded for proteins of 18-50.5 kDa, with a pI range of 5.4-7.8. These new stress mRNAs represented one of two main classes. Class I consisted of mRNAs shared by both genotypes. Class II represented mRNAs specific to each genotype; unique mRNAs of roots accumulated preferentially in the salt-tolerant genotype, whereas those of shoots accumulated in the salt-sensitive genotype. The findings suggest that transcriptional as well as posttranscriptional mechanisms regulate gene expression in barley during salinity stress.

Journal Article↗

Arbuscular mycorrhizal relations of mangrove plant community at the Ganges river estuary in India.

Mangroves are climax formation of hydrohalophytes inhabiting estuarine or marine salt marshes in the tropics and subtropics. As a terrestrial plant community inhabiting tidally inundated estuarine or marine sediments, mangroves show considerable adaptation to salinity, water-logging and nutrient stress. Thirty-one species of mangrove and mangrove associates and 23 species of transported flora, belonging to 25 families at four physiographic stages of succession of the mangrove plant community at the terminal part of the Ganges river estuary in India were examined for arbuscular mycorrhizal (AM) root association. Dominant members of the mangrove plant community were all AM, mostly with 'Paris' type structures. Many of the known non-mycotrophic plant families, except the Cyperaceae, also showed AM association, with intracellular hyphae and vesicles as the most discernible endophyte structures. Intensity of AM colonization varied both with the species and situations of their occurrence, being more intense and also more extensive in less saline dry ridge mangroves than in more saline formative and developed swamp mangroves. Introduced exotic trees on the ridges and embankments were infected by AM, but less than the declining mangroves in the same location. Seven species of AM fungi in common with those of the upstream mesophytic plants were isolated from root-free rhizosphere soils of the mangroves, three of which predominated in root association. These species, individually and as mixtures, infected roots of salinity tolerant herbs and trees in both locational silt and upstream alluvial soil with obvious improvements in their biomass yield and phosphorus nutrition. AM infective potential of root-free rhizosphere soils of the dominant members of the mangrove community were negatively related to salinity level of the sediment soil of the successional stages. The evidences of AM association of mangroves and other salt marsh plants obtained here and those reported elsewhere are discussed.

Avicennia↗

Does exogenous application of salicylic acid through the rooting medium modulate growth and photosynthetic capacity in two differently adapted spring wheat cultivars under salt stress?

In order to assess whether exogenous application of salicylic acid (SA) through the rooting medium could modulate the photosynthetic capacity of two wheat cultivars differing in salinity tolerance, a hydroponic experiment was conducted under greenhouse conditions. Seeds of a salt tolerant (S-24) and a moderately salt sensitive (MH-97) cultivar were germinated at 0 or 150 mM NaCl in Hoagland's nutrient solution containing different levels of salicylic acid (SA) (0, 0.25, 0.50, 0.75 and 1.00 mM) for 7d. Seven-day old wheat seedlings were transferred to hydroponics and grown at 0, or 150 mM NaCl for for further 30 d. Different levels of salicylic acid (SA) were also maintained in the solution culture. After 30 d, four plants out of six were harvested and the remaining plants were left for the estimation of yield attributes Salt stress reduced the growth and grain yield of both cultivars. However, cv. S-24 performed better than MH-97 under salt stress with respect to leaf area, and grain yield. Exogenous application of SA promoted growth and yield, and counteracted the salt stress-induced growth inhibition of salt tolerant S-24, whereas for MH-97 there was no improvement in growth or grain yield with SA application. Of the varying SA levels used, the most effective levels for promoting growth and grain yield were 0.75 and 0.25 mM under normal and saline conditions, respectively. The improvement in growth and grain yield of S-24 due to SA application was associated with improved photosynthetic capacity. Changes in photosynthetic rate due to SA application were not due to stomatal limitations, but were associated with metabolic factors, other than photosynthetic pigments and leaf carotenoids.

Adaptation, Physiological↗