Arguments for the use of physiological criteria for improving the salt tolerance in crops

Efforts to develop new crop varieties with improved salt tolerance have been intensified over the past 15–20 years. Despite the existence of genetic variation for salt tolerance within species, and many methods available for expanding the source of genetic variation, there is only a limited number of varieties that have been developed with improved tolerance. These new varieties have all been based upon selection for agronomic characters such as yield or survival in saline conditions. That is, based upon characters that integrate the various physiological mechanisms responsible for tolerance. Yet over the same time period, knowledge of physiological salt responses has increased substantially.Selection and breeding to increase salt tolerance might be more successful if selection is based directly on the physiological mechanisms or characters conferring tolerance. Basic questions associated with using physiological selection criteria are discussed in the paper. These are centred around the need for genetic variation, the importance of the targeted mechanism, the ease of detection of the physiological mechanism (including the analytical requirements) and the breeding strategy. Many mechanisms, including ion exclusion, ion accumulation, compatible solute production and osmotic adjustment have been associated with genetic variation in salt tolerance. Yet their successful use in improving salt tolerance, via physiological selection criteria, is largely non-existent. Consideration is given to the role of physiological criteria in the short and long term in improving salt tolerance. In several glycophytic species, particularly legumes, physiological selection based on ion exclusion from the shoots shows promise. Recent results for white clover indicate the potential for using a broad physiological selection criterion of restricted Cl accumulation in the shoots, with scope for future refinement based upon the specific physiological characters that combined result in ion exclusion.     

Effect of aluminium on the growth of 34 plant species: A summary of results obtained in low ionic strength solution culture

The results from many experiments conducted over 5 years to determine the tolerance of 34 plant species (87 cultivars) to aluminium (Al) are summarised. All experiments were conducted in a temperature-controlled glasshouse using a low-ionic-strength solution culture technique. The activity of Al³⁺ (M) at which top yields were reduced by 50% (Al_RY50) was determined for each cultivar.The species Bromus wildenowii, Cynosurus cristatus, Hordeum vulgare, Triticum aestivum (cvs Warigal, Scout, Sonora-63), Avena byzantina, Arabidopsis thaliana, Lycopersicon esculentum and Nicotiana plumbaginifolia were all very sensitive to Al (Al_RY50<1). The species Poa pratense, Lolium perenne (NZ-derived cultivars), Lotus corniculatus, Avena sativa (cvs West, Carbeen, Camellia and Coolabah), Triticum aestivum (cvs Cardinal and Waalt), Allium cepa and Asparagus officinalis were sensitive to Al (Al_RY50 1–2).The pasture grass species Lolium perenne (Australian and European and derived cultivars), Lolium hybridum and Lolium multiflorum, Dactylis glomerata (Apanui and Kara), Phalaris aquatica, Festuca arundinacea and the pasture legumes species Trifolium pratense, Trifolium repens and Trifolium subterraneum were all moderately sensitive to Al (Al_RY50 2–5). Other species that were also moderately sensitive included Triticum aestivum (cvs Atlas-66, BH146, and Carazinho), Avena sativa (cvs Swan and Blackbutt), Avena Strigosa, Petunia x and Phaseolus vulgaris (cvs Red Kidney, Black Turtle and Haricot).The most tolerant species (Al_RY50>5) were (in order of increasing tolerance) Phaseolus vulgaris (cvs Tendergreen, The Prince and Yatescrop), Cucurbita maxima, Dactylis glomerata (cv Wana), Paspalum dilatatum, Lotus pedunculatus, Ehrharta calycina, Medicago sativa, Holcus lanatus, Festuca rubra, Phaseolus lunatus and Agrostis tenuis. Agrostis tenuis was at least twice as tolerant as the next most tolerant species (Al_RY50>30 compared to 15.6).     

Measuring maximum root growth instead of longest root elongation in metal tolerance tests for grasses (Agrostis capillaris,Agrostis delicatula and Agrostis castellana)

Longest root elongation diminished significantly in the three species tested from 6 mm d^-1 to 3 mm d^-1 in 3 weeks. During this period S.D. increased considerably (from 49% to 112%, A. castellana), and accounted on the average for 68% (A. capillaris) till 94% (A. castellana) of the mean. Maximum root growth stabilized at 6 mm d^-1 and showed less variation in the measurements (S.D. 52% of the mean). Growth of the originally longest root approaches zero in all three species, in accordance with the natural cease of growth of roots in grasses fascicular root system. Measuring maximum root growth instead of longest root elongation is proposed for testing metal tolerance of grasses in sequential experiments.     

The effects of excess manganese on photosynthetic rate and concentration of chlorophyll in Triticum aestivum grown in solution culture

Manganese toxicity, which involves a broad array of physiological responses, has been identified as an important factor limiting plant growth on acid soils. In the experiments reported here, we examined the toxic effects of Mn on chlorophyll content, photosynthesis and respiration in two cultivars (Norquay and Columbus) of Triticum aestivum (wheat) which differ in tolerance of Mn. When grown over a range of concentrations of Mn (0–1 000 μ M ), the Mn-tolerant cultivar maintained higher rates of photosynthesis and respiration, and higher concentrations of chlorophyll a and chlorophyll b , than did the Mn-sensitive cultivar, despite greater accumulations of Mn in leaf tissues. After 5 days growth with 1 000 μ M Mn in solution, the photosynthetic rate fell to 25% of control in the sensitive cultivar and to only 75% of control in the tolerant cultivar. The concentration of chlorophyll a fell to 50% of control in the sensitive cultivar, but did not differ from control in the tolerant cultivar. Greater effects were seen on concentrations of chlorophyll b . which fell to 35% and 55% of control in the sensitive and tolerant cultivars, respectively. Rates of photosynthesis decreased in both cultivars as concentrations of chlorophyll decreased; however, the photosynthetic rate per unit chlorophyll remained constant or increased in the tolerant cultivar and decreased in the sensitive cultivar as concentrations of Mn in solution increased. Thus, in the sensitive cv. Columbus, Mn seemed to have a toxic effect on both chlorophyll content and photosynthesis per unit chlorophyll. In the tolerant cv. Norquay, the only clear effect of Mn was a reduction in chlorophyll content, although direct inhibition of photosynthesis could not be discounted.     

Drought tolerance in faster- and slower-growing black spruce (Picea mariana) progenies: II. Osmotic adjustment and changes of soluble carbohydrates and amino acids under osmotic stress

The possibility was considered that osmotic adjustment, the ability to accumulate solutes in response to water stress, may contribute to growth rate differences among closely-related genotypes of trees. Progeny variation in osmotic adjustment and turgor regulation was investigated by comparing changes in osmotic and pressure potentials, soluble carbohydrates, and amino acids in osmotically stressed seedlings in 4 full-sib progenies of black spruce [ Picea mariana (Mill.) B. S. P.] that differed in growth rate under drought. Osmotic stress was induced by a stepwise increase in the concentration of polyethylene glycol (PEG)-3350 from 10 (w/v) to 18 and 25%, which provided osmotic potentials in solution culture of -0.4, -1.0 and -2.0 MPa each for 3 days. All 4 progenies maintained a positive cell turgor even at 25% PEG, due to a significant decline in osmotic potential. Although total amino acids, principally proline, increased, ca 60% of the decrease in osmotic potential was attributable to soluble carbohydrates and glucose was the major osmoregulating solute. There was little progeny variation in any of measured parameters in unstressed seedlings. Compared to two slower-growing progenies, the two progenies capable of more vigorous growth under drought in the field accumulated more soluble carbohydrates (mainly glucose and fructose), developed lower osmotic potential and maintained higher turgor pressure when osmotically-stressed in solution culture. The ability to adjust osmotically and maintain turgor under drought stress could thus be a useful criterion for the early selection of faster-growing, drought-tolerant genotypes.     

Root-zone temperature and salinity: Interacting effects on tillering,growth and element concentration in barley

The effects of root-zone salinity (0, 30, and 60 mmol L^–1 of NaCl) and root-zone temperature (10, 15, 20, and 25°C) and their interactions on the number of tillers, total dry matter production, and the concentration of nutrients in the roots and tops of barley (Hordeum vulgare L.) were studied. Experiments were conducted in growth chambers (day/night photoperiod of 16/8 h and constant air temperature of 20°C) and under water-culture conditions. Salinity and root temperature affected all the parameters tested. Interactions between salinity and temperature were significant (p<0.05) for the number of tillers, growth of tops and roots, and the concentration of Na, K, P in the tops and the concentration of P in the roots. Maximum number of tillers and the highest dry matter were produced when the root temperature was at the intermediate levels of 15 to 20°C. Effect of salinity on most parameters tested strongly depended on the prevailing root temperature. For example, at root temperature of 10°C addition of 30 mmol L^–1 NaCl to the nutrient solution stimulated the growth of barley roots; at root temperature of 25°C, however, the same NaCl concentration inhibited the root growth. At 60 mmol L^–1, root and shoot growth were maximum when root temperature was kept at the intermediate level of 15°C; most inhibition of salinity occurred at both low (10°C) and high (25°C) root temperatures. As the root temperature was raised from 10 to 25°C, the concentration of Na generally decreased in the tops and increased in the roots. At a given Na concentration in the tops or in the roots, respective growth of tops or roots was much less inhibited if the roots were grown at 15–20°C. It is concluded that the tolerance of barley plant to NaCl salinity of the rooting media appears to be altered by the root temperature and is highest if the root temperature is kept at 15 to 20°C.     

Heavy metal tolerance in Festuca ovina L. from contaminated sites in the Eifel Mountains,Germany

Extremely high degrees of lead tolerance, measured by comparing rates of root extension in culture solutions, are reported from populations of Festuca ovina growing at two lead-mining sites (Westschacht and Keldenich-II) near Mechernich in the Eifel Mountains, Germany. Other populations from nearby heavy metal-contaminated areas show a considerably smaller degree of lead tolerance. Samples of Festuca ovina collected in the field at Westschacht and Keldenich-II contain higher levels of lead in their aerial organs than do those from other lead-contaminated sites. The main soil factor determining the high degree of lead tolerance is the high Pb/Ca ratio. Populations from soils with a low Pb/Ca ratio display a very low degree of tolerance. It is therefore concluded that in Westschacht and Keldenich-II plants, a genuine intracellular tolerance mechanism is present, allowing the accumulation of lead in aerial organs.Leaf samples of zinc-tolerant Festuca contain higher levels of zinc than do samples of non-tolerant plants. Lead and zinc amounts in leaves are correlated with the soil ratios of Pb/Ca and Zn/Ca, respectively, rather than with the absolute soil-metal levels.In a slightly lead-tolerant, but highly zinc-tolerant clone of Festuca ovina from a site contaminated with large amounts of lead and zinc (Plombières), lead was found to be the major factor affecting the inhibition of root extension with combined treatments of lead and zinc in culture solutions. In the highly lead-tolerant, zinc-sensitive population from Westschacht, zinc governs the response of root growth to combinations of the two metals. The results are discussed in terms of discriminating distinct types of heavy-metal tolerance.     

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