Excretory organ growth and implications for salt tolerance in hatchling American avocets Recurvirostra americana

The role of salt glands in avian osmoregulation has been widely studied. Acclimation to saline habitats in aquatic birds involves increases in the relative size and complexity of the salt glands, and it is generally agreed that salt gland size varies as a function of salt loading, and is broadly correlated with habitat salinity. We report here salt gland sizes in three age classes of American avocet Recurvirostra americana chicks. Mean relative (mg/g body mass) salt gland masses for newly hatched (<24 h old) avocet chicks collected at a wetland supplied with pumped fresh water was 0.48 mg salt gland/g body mass. This value is comparable to the low end of published values for some strictly marine birds and similar to values for adult American avocets collected at saline lakes. These results suggest that American avocets, which are frequently raised in saline environments, hatch with salt glands that are large enough to cope with the osmoregulatory demands of saline environments.     

The American Avocet (Recurvirostra americana) as a paradigm for adult automimicry

Summary The body of theory concerning life-history strategies predicts that the duration of high-mortality stages should be minimized by natural selection. This is especially applicable to the avian pre-flight stage, during which growth rates typically are rapid. Using the American Avocet (Recurvirostra americana) as a paradigm, I propose a developmental strategy by which young animals can lower their mortality rates by an accelerated (and deceptive) acquisition of adult or adult-like characters. The benefit accrues when predators misidentify the vulnerable young as adults and fail to attack them because adults are much less vulnerable. This strategy, termed adult automimicry, is most likely to occur in precocial species living in open habitats.American Avocets are large, precocial, open-country shorebirds that first fly when about 4–5 weeks old. They develop a juvenal, plumage in their third week that resembles adult breeding plumage in pattern and color, even though plumage details are different. At this time chicks begin using adult foraging techniques and tend to move away rather than hide from potential predators. A few weeks later they acquire a first winter plumage that resembles adult winter plumage. Thus, avocet chicks appear unusually adult-like after their second week. This should make it difficult for distant predators to distinguish flightless chicks from volant adults.     

Breeding tomatoes for salt tolerance: inheritance of salt tolerance and related traits in interspecific populations

Summary Interspecific segregating populations derived from a cross between tomato (Lycopersicon esculentum) cv M82-1 -8 (M82) and the wild species L. pennellii accession LA-716 (Lpen716) were used to study the genetic basis of salt tolerance and its implications for breeding. BC₁ (M82 x (M82 x Lpen716)) and BC₁ S₁ (progenies of selfed BC₁ plants) populations were grown under arid field conditions and irrigated with water having electrical conductivities of 1.5 (control), 10 and 20 dSm^-1. The evaluation of salt tolerance was based on total fruit yield (TY), total dry matter (TD) and TD under salinity relative to the control (RD). Sodium, potassium and chloride concentrations were measured in the leaves and stems. The methods for estimating heritability were adapted to BC₁ plants and BC₁S₁ families. TY, TD and RD had heritability estimates of 0.3–0.45, indicating that salt tolerance can be improved by selection. Genetic correlations between traits indicated that high yield may be combined with salt tolerance and that ion contents are not likely to provide an efficient selection criteria for salt tolerance. Genetic correlations between performances under various salinity levels suggested that similar mechanisms affect the responses to salinity treatments of 10 and 20 dSm^-1. Responses to paper selection confirmed that salt tolerance of the tomato may be improved by selection, and that this selection should be based on dry matter and yield parameters under salinity.Passed away May 1986     

Advances in salt tolerance

Summary Advances in and prospects for the development of salt tolerant crops are discussed. The genetic approach to the salinity problem is fairly new, but research has become quite active in a short span of time. Difficulties and opportunities are outlined. Salinity varies spatially, temporally, qualitatively, and quantitatively. In addition, the responses of plants to salt stress vary during their life cycle. Selection and breeding, including the use of wide crosses, are considered the best short-term approaches to the development of salt tolerant crops, but the new biotechnological and molecular biological techniques will make increasingly important contributions. Cooperation is called for among soil and water scientists, agronomists, plant physiologists and biochemists, cytologists, and plant geneticists, breeders, and biotechnologists. Given such cooperation and adequate support for these endeavors, the potential for increasing productivity in salt-affected areas can be realized.     

Principles and strategies in breeding for higher salt tolerance

Summary Salinity is an environmental component that usually reduces yield. Recent advances in the understanding of salt effects on plants have not revealed a reliable physiological or biochemical marker that can be used to rapidly screen for salt tolerance. The necessity of measuring salt tolerance based upon growth in saline relative to non-saline environments makes salt tolerance measurements and selection for tolerance difficult. Additionally, high variability in soil salinity and environmental interactions makes it questionable whether breeding should be conducted for tolerance or for high yield. Genetic techniques can be used to identify the components of variation attributable to genotype and environment, and the extent of genetic variation in saline and nonsaline environments can be used to estimate the potential for improving salt tolerance. Absolute salt tolerance can be improved best by increasing both absolute yield and relative salt tolerance.     

Identification and validation of QTLs for salt tolerance during vegetative growth in tomato by selective genotyping.

The purpose of this study was to identify quantitative trait loci (QTLs) for salt tolerance (ST) during vegetative growth (VG) in tomato by distributional extreme analysis and compare them with the QTLs previously identified for this trait. A BC1 population (N = 792) of a cross between a moderately salt-sensitive Lycopersicon esculentum Mill. breeding line (NC84173, maternal and recurrent parent) and a salt-tolerant L. pimpinellifolium (Jusl.) Mill. accession (LA722) was evaluated for ST in solution cultures containing 700 mM NaCl + 70 mM CaCl2 (electrical conductivity, EC = 64 dS/m and phiw approximately -35.2 bars). Thirty-seven BC1 plants (4.7% of the total) that exhibited the highest ST were selected (referred to as the selected population), grown to maturity in greenhouse pots and self-pollinated to produce BC1S1 progeny seeds. The 37 selected BC1S1 progeny families were evaluated for ST and their average performance was compared with that of the parental BC1 population before selection. A realized heritability of 0.50 was obtained for ST in this population. The 37 selected BC1 plants were subjected to restriction fragment length polymorphism (RFLP) analysis using 115 markers, and marker allele frequencies were determined. Allele frequencies for the same markers were also determined in an unselected BC1 population (N = 119) of the same cross. A trait-based marker analysis (TBA), which measures differences in marker allele frequencies between selected and unselected populations, was used to identify marker-linked QTLs. Five genomic regions were detected on chromosomes 1, 3, 5, 6, and 11 bearing significant QTLs for ST. Except for the QTL on chromosome 3, all QTLs had positive alleles contributed from the salt tolerant parent LA722. Of the five QTLs, three (those on chromosomes 1, 3, and 5) were previously identified for this trait in another study, and thus were validated here. Only one of the major QTLs that was identified in our previous study was not detected here. This high level of conformity between the results of the two studies indicates the genuine nature of the identified QTLs and their potential usefulness for ST breeding using marker-assisted selection (MAS). A few BC1S1 families were identified with most or all of the QTLs and with a ST comparable to that of LA722. These families should be useful for the development of salt tolerant tomato lines via MAS.     

Increasing salt tolerance in the tomato

In this paper, a number of strategies to overcome the deleterious effects of salinity on plants will be reviewed; these strategies include using molecular markers and genetic transformation as tools to develop salinity-tolerant genotypes, and some cultural techniques. For more than 12 years, QTL analysis has been attempted in order to understand the genetics of salt tolerance and to deal with component traits in breeding programmes. Despite innovations like better marker systems and improved genetic mapping strategies, the success of marker-assisted selection has been very limited because, in part, of inadequate experimental design. Since salinity is variable in time and space, experimental design must allow the study of genotype x environment interaction. Genetic transformation could become a powerful tool in plant breeding, but the growing knowledge from plant physiology must be integrated with molecular breeding techniques. It has been shown that the expression of several transgenes promotes a higher level of salt tolerance in some species. Despite this promising result, the development of a salt-tolerant cultivar by way of transgenesis has still not been achieved. Future directions in order to overcome the present limitations are proposed. Three cultural techniques have proved useful in tomato to overcome, in part, the effects of salinity: treatment of seedlings with drought or NaCl ameliorates the adaptation of adult plants to salinity; mist applied to tomato plants grown in Mediterranean conditions improves vegetative growth and yield in saline conditions; and grafting tomato cultivars onto appropriate rootstocks could reduce the effects of salinity.     

Vegetative salt tolerance of barnyard grass mutants selected for salt tolerant germination

Improving salt tolerance of economically important plants is imperative to cope with the increasing soil salinity in many parts of the world. Mutation breeding has been widely used to improve plant performance under salinity stress. In this study, we have mutagenized Echinochloa crusgalli L. with sodium azide and three selected mutants (designated fows A) with salt tolerant germination. Their vegetative growth was compared to that of the wild type after short-term and long-term salt stress. The germination of the three fows A mutants in the presence of inhibitory concentrations of NaCl, KCL, and mannitol was better than that of the wild type. Early growth of the mutants in the presence of 200 mM NaCl was also better than that of the wild type perhaps due to improved K⁺ uptake and enhanced accumulation of sugars particularly sucrose at least in two mutants. But the three mutants and the wild type responded similarly to long-term salt stress. The tolerance mechanisms during short-term and long-term salt stress are discussed.     

Co-existence of salt and drought tolerance in Triticeae

Cell membrane stability (CMS) technique was used to screen for drought tolerance, salt tolerant accessions of three Aegilops species, Ae. tauschii, Ae. cylindrica, Ae. geniculata and two hexaploid wheat (Tricitum aestivum L.) cultivars comprising salt tolerant LU-26 and drought tolerant Chakwal-86. The objectives were to see how valid it is for a salt tolerant plant to be drought tolerant as well and to identify the character(s) that may contribute to drought tolerance. Three moisture levels equal to 100, 50 and 25% saturation capacity of the soil were used for plant cultivation. Injury percentage (IP) based on in-vitro desiccation induced by polyethylene glycol (PEG) in leaf tissue was measured through the conductivity of the electrolyte leakage. Injury percentage decreased in all the test material with decrease in soil moisture contents. Ae. cylindrica exhibited minimum injury at 100% soil moisture level followed by Ae. tauschii and Ae. geniculata while drought tolerant wheat cultivars exhibited the maximum. The wheat cultivar Chakwal-86 has been developed for dry areas, with low soil moisture levels, and high water potential enhances the injury percentage. Aegilops cylindrica is a salt tolerant species and can thus tolerate water deficit conditions created due to low osmotic potential. Potassium appeared to play an important role in drought tolerance which was evident from high K+ contents and low K+ leakage from Aegilops cylindrica and drought tolerant wheat cultivar Chakwal-86. It was inferred from the study that salt tolerant species might prove drought tolerant in the areas where water deficit prevails due to the ability to create low intracellular osmotic potentials.     

Some important physiological selection criteria for salt tolerance in plants

Undoubtedly, plant breeders have made a significant achievement in the past few years, improving salinity tolerance in a number of potential crops using artificial selection and conventional breeding approaches, although molecular biology approaches are currently being intensively pursued for achieving this goal. However, most of the selection procedures used so far, were based merely on differences in agronomic characters. Agronomic characters represent the combined genetic and environmental effects on plant growth, and include the integration of the physiological phenomena conferring salinity tolerance. In fact, physiological criteria are able to supply more reliable information than agronomic characters. Although there are large numbers of reports in the literature mainly dealing with water relations, photosynthesis, and accumulation of various inorganic ions and organic metabolites in individual crops, there is little information available on the use of these attributes as selection criteria for improving salt tolerance through selection and breeding programs. In this review, the major adaptive components of salt tolerance such as osmotic adjustment, photosynthesis, water relations and ion relations are reviewed. In view of the complexity of salt tolerance and its great variation at intra-specific and inter-specific levels, it is difficult to identify single criteria, which could be used as effective selection targets. Rather it is most meaningful if physiological and biochemical indicators for individual species are determined rather than generic indicators.     

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.     

Control of salt gland activity in the hatchling green sea turtle, Chelonia mydas

 We studied the control of salt gland secretion in hatchling Chelonia mydas. The threshold salt load to activate salt secretion was between 400 μmol NaCl 100 g bodymass (BM)⁻¹ and 600 μmol NaCl 100 g BM⁻¹, which caused an increase in plasma sodium concentration of 13% to 19%. Following a salt load of 2700 μmol NaCl 100 g BM⁻¹, salt gland secretion commenced in 12 ± 1.3 min and reached maximal secretory concentration within 2–7 min. Maximal secretory rate of a single gland averaged 415 μmol Na 100 g BM⁻¹ h⁻¹. Plasma sodium concentration and total osmotic concentration after salt loading were significantly higher than pretreatment values within 2 min. Adrenalin (25 μg kg BM⁻¹) and the cholinergic agonist methacholine (1 mg kg BM⁻¹) inhibited salt gland activity. Atropine (10 mg kg BM⁻¹) reversed methacholine inhibition and stimulated salt gland secretion when administered with a subthreshold salt load. Arginine vasotocin produced a transient reduction in sodium secretion by the active gland, while atrial natriuretic factor, vasoactive intestinal peptide and neuropeptide Y had no measurable effect on any aspect of salt gland secretion. Our results demonstrated that secretion of the salt gland in C. mydas can be modified by neural and hormonal chemicals in vivo and that the cholinergic and adrenergic stimulation of an exocrine gland do not appear to have the typical, antagonist actions on the chelonian salt gland. Accepted: 28 September 1999     

Dispersal and genetic structure in the American marten, Martes americana

Natal dispersal in a vagile carnivore, the American marten (Martes americana), was studied by comparing radio-tracking data and microsatellite genetic structure in two populations occupying contrasting habitats. The genetic differentiation determined among groups of individuals using F(ST) indices appeared to be weak in both landscapes, and showed no increase with geographical distance. Genetic structure investigated using pairwise genetic distances between individuals conversely showed a pattern of isolation by distance (IBD), but only in the population occurring in a homogeneous high-quality habitat, therefore showing the advantage of individual-based analyses in detecting within-population processes and local landscape effects. The telemetry study of juveniles revealed a leptokurtic distribution of dispersal distances in both populations, and estimates of the mean squared parent-offspring axial distance (sigma2) inferred both from the genetic pattern of IBD and from the radio-tracking survey showed that most juveniles make little contribution to gene flow.     

Autophagy is required for tolerance of drought and salt stress in plants

Autophagy is a protein degradation process in which cells recycle cytoplasmic contents when subjected to environmental stress conditions or during certain stages of development. Upon the induction of autophagy, a double membrane autophagosome forms around cytoplasmic components and delivers them to the vacuole or lysosome for degradation. In plants, autophagy has been shown previously to be induced during abiotic stresses including nutrient starvation and oxidative stress. In this paper, we demonstrate the induction of autophagy in high salt and osmotic stress conditions, concomitant with the upregulation of expression of an Arabidopsis thaliana autophagy-related gene AtATG18a. Autophagy-defective RNAi-AtATG18a plants are more sensitive to salt and drought conditions than wild-type plants, demonstrating a role for autophagy in the response to these stresses. NADPH oxidase inhibitors block autophagy induction upon nutrient starvation and salt stress, but not during osmotic stress, indicating that autophagy can be activated by NADPH oxidase-dependent or -independent pathways. Together our results indicate that diverse environmental stresses can induce autophagy and that autophagy is regulated by distinct signaling pathways in different conditions.