Abstract

Comparative studies on the responses to salt stress of taxonomically related taxa should help to elucidate relevant mechanisms of stress tolerance in plants. We have applied this strategy to three Plantago species adapted to different natural habitats, P. crassifolia and P. coronopus–both halophytes–and P. major, considered as salt-sensitive since it is never found in natural saline habitats. Growth inhibition measurements in controlled salt treatments indicated, however, that P. major is quite resistant to salt stress, although less than its halophytic congeners. The contents of monovalent ions and specific osmolytes were determined in plant leaves after four-week salt treatments. Salt-treated plants of the three taxa accumulated Na+ and Cl- in response to increasing external NaCl concentrations, to a lesser extent in P. major than in the halophytes; the latter species also showed higher ion contents in the non-stressed plants. In the halophytes, K+ concentration decreased at moderate salinity levels, to increase again under high salt conditions, whereas in P. major K+ contents were reduced only above 400 mM NaCl. Sorbitol contents augmented in all plants, roughly in parallel with increasing salinity, but the relative increments and the absolute values reached did not differ much in the three taxa. On the contrary, a strong (relative) accumulation of proline in response to high salt concentrations (600–800 mM NaCl) was observed in the halophytes, but not in P. major. These results indicate that the responses to salt stress triggered specifically in the halophytes, and therefore the most relevant for tolerance in the genus Plantago are: a higher efficiency in the transport of toxic ions to the leaves, the capacity to use inorganic ions as osmotica, even under low salinity conditions, and the activation, in response to very high salt concentrations, of proline accumulation and K+ transport to the leaves of the plants.

Highlights

  • Drought and soil salinity belong to the environmental factors most adverse for plants, which cause the biggest losses in agricultural production throughout the world and determine to a PLOS ONE | DOI:10.1371/journal.pone.0160236 August 4, 2016large extent the distribution of wild species in nature [1, 2]

  • The average number of leaves per plant varied from 12 to 45 for control, nonstressed plants, while at the highest salt concentration tested (800 mM NaCl) this number was reduced by 43% in P. coronopus, by 51% in P. crassifolia and by 54% in P. major (Fig 1)

  • A similar pattern of reduction in the number of leaves, in relation to the non-treated controls, was observed in the plants that survived the eight-week salt treatments, up to 400 mM NaCl. According to this criterion, P. coronopus appears to be the most salt-tolerant of the three species, which is supported by the fact that a significant decrease in the mean leaf number was only observed in this species at high salt concentrations

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Summary

Introduction

Drought and soil salinity belong to the environmental factors most adverse for plants, which cause the biggest losses in agricultural production throughout the world and determine to a PLOS ONE | DOI:10.1371/journal.pone.0160236 August 4, 2016large extent the distribution of wild species in nature [1, 2]. The vast majority of plants, including all major crops, are glycophytes (that is, salt sensitive), there are some species naturally adapted to saline environments, named halophytes. They represent only about 0.25% of total angiosperm taxa, but are very diverse from a taxonomic point of view, as tolerance to salinity seems to have evolved independently in different plant lineages [3]. Stress tolerance in plants relies on the activation of a series of conserved response pathways, some of them common to different abiotic stresses One of these basic response mechanisms is the control of ion homeostasis and maintenance of osmotic balance, to counteract cellular dehydration caused by soil salinity, drought, cold or high temperatures, among other stressful conditions. The relative resistance to salt stress of plants, which varies widely among taxa, should be attributed to quantitative rather than qualitative differences in their mechanisms of response, which only in halophytes are efficient enough to confer salt tolerance, always within species-specific limits [7,8,9,10]

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