Desiccation tolerance (DT) of seeds, one of the plant's environmental adaptation mechanisms, allows them to survive as seeds in a quiescent state under extremely water-deficient conditions during the plant's life cycle, followed by seed germination and seedling establishment under favourable water conditions. The seed-DT is lost after radicle emergence; however, there is a developmental period called the ‘DT window’ during which the germinating seeds can re-induce DT following a cue from their ambient low water potential (i.e. mild osmotic stress). The DT re-inducibility within the DT window has been used as a model biosystem for understanding molecular mechanisms that activate/supress DT in a number of plant species. However, the characteristics of the DT window for species particularly important to the agroindustry are still largely fragmented. Here, physiological analyses were performed, aiming to elucidate the properties of the DT window in tomato, a model species for Solanaceae, holding a key strategic position for the seed industry and commercial use around the world. We revealed that (i) the DT window of tomato seeds is closed when the developing radicle reaches about 4 mm after germination, (ii) the most effective ambient water potential to re-induce DT into seeds is about −1.5 MPa and (iii) there is organ specificity of DT re-induction with hypocotyls, showing a longer DT window than cotyledons and roots in post-germination seeds.

We investigated the thermal thresholds to seed germination and the variations in abscisic acid (ABA) levels and oxidative metabolism during seed dormancy-breaking and germination in two palm species with differences in desiccation tolerance. We used Mauritia flexuosa (buriti palm, desiccation-sensitive seeds) from swampy habitats (Veredas) and Attalea speciosa (babassu, desiccation-tolerant seeds) from the transition zone between the forest and semi-arid region (drained soils). Germination was evaluated at 15–40°C after dormancy-breaking (operculum removal). At optimal temperature for both species (30°C), embryos were sampled in distinct germination phases – dry, imbibed, after operculum removal and at early germination – and used for quantifying ABA and hydrogen peroxide (H2O2) content, antioxidant enzyme activities and for histolocalization of superoxide anion (O2−). Seeds of M. flexuosa germinated only in a narrow temperature range (25–35°C), while A. speciosa seeds germinated between 15 and 40°C. After operculum removal, reduced ABA levels in embryos of M. flexuosa were accompanied by constant H2O2 levels, while in A. speciosa, similar levels of ABA and H2O2 were maintained throughout all germination phases. The presence of O2− was restricted to the haustorium, and an increase in O2− accumulation was observed in both species after operculum removal. Similarities were noted between both species regarding enzyme activities; however, the activities were higher in embryos from M. flexuosa. The presence of O2− only in the haustorium indicates that this region of the embryo is an active structure following imbibition and is involved in germination itself, not just functioning in reserve mobilization.

Lichen extinction occurs at rapid rates as a result of human activity, although species could potentially be rescued by conservation management based on ecophysiological knowledge. The boreal old forest cyanolichen Erioderma pedicellatum currently occurs in few sites worldwide. To protect it from extinction, it is essential to learn more about it. The last remaining good European site is a canyon with a waterfall, in a low-rainfall region of Norway. Here, a spatially restricted population of 1500–2000 thalli dominates the epiphytic vegetation of a small number of Picea abies canopies. We were able to document that 1) E. pedicellatum grew on thin branches with higher bark pH than is normal for P. abies in a canyon that provided an unusual combination of very high light, high air humidity, and cool temperatures in the growing season. However, the species did not inhabit the main waterfall spray zone. 2) Erioderma pedicellatum had a high light saturation point, high CO2 uptake at high light (≥ 600 μmol m−2 s−1) and cool temperatures (5–20 °C), and experienced strong suprasaturation depression of photosynthesis when fully hydrated. 3) It showed good tolerance of desiccation and high light; it was slightly more tolerant than the morphologically similar, but more common cyanolichen Pectenia plumbea. 4) The European population in its sunny habitat had higher water holding capacity than previously recorded in slightly shaded rainforest populations in Newfoundland, consistent with acclimation to compensate for high evaporative demands. Understanding the ecological niche and responses to critical environmental factors is essential for action plans to avoid extinction of E. pedicellatum. Methods used in this study could also be applicable for ecological understanding of other threatened lichen species.

Functional traits related to regeneration responses to the environment are highly determinants of distribution patterns of plant communities. A large body of studies on seed traits suggests that regional climate may act as a strong filter of plant recruitment; however, few studies have evaluated the relative importance of seed traits and environmental filters for seed persistence at the population level. We tested the role of seed mass, water content and desiccation tolerance, as well as the germination time as proxies for seed tolerance to environmental filters (water deficit, heat shock and high temperatures) by comparing the response of tree species co-occurring in savannas located in different regions: Cerrado biome of Central Brazil and the Rio Branco savannas of northern Brazil. Seeds collected in savannas of Rio Branco showed a higher tolerance to environmental filters than those collected in savannas of the Cerrado. While the germination percentages largely varied in response to the treatments, the germination times were virtually unaffected by them, irrespective of seed origin, seed mass and water content. At the population level, the regional environment was a key determinant of seed tolerance to stress, irrespective of seed traits. Germination time was shown to represent a conservative seed trait and more linked to a species-specific germination strategy than to regional characteristics. Our results suggest that recruitment patterns of Cerrado savannas may be more impacted than Rio Branco savannas by the climate scenarios predicted for the future.

The astonishing capability of life to adapt to extreme conditions has provided a new perspective on what ‘habitable’ means. On Earth extremophiles thrive in hostile habitats, such as hot and cold deserts or Antarctic sub-glacial lakes considered as Earth analogues of Mars and icy moons of Jupiter and Saturn. Recently desert cyanobacteria were exposed to ground-based simulations of space and Martian conditions and to real space and Martian conditions simulated in low Earth orbit using facilities attached outside the International Space Station. When exposure to such conditions does not exceed repair capabilities, more data are available regarding the physico-chemical constraints that life can withstand. When the accumulated damage exceeds the survival potential, the persistence of biomarkers contributes to the search for life elsewhere. Knowledge concerning the endurance of desert cyanobacteria under space and Martian conditions contributes to the development of life support systems.

Solute leakage is used as an indicator of membrane damage during desiccation of recalcitrant seeds. We re-examined this phenomenon and its utility by comparing recalcitrant Spartina alterniflora and orthodox Spartina pectinata seeds, as well as drying isolated embryos or embryos isolated after whole-seed drying. During drying, intact seed leakage (electrical conductivity and absorbance at 280 nm) was independent of seed moisture content, dormant/non-dormant state or desiccation tolerance. Embryos from both Spartina species, isolated after drying within the intact seed, exhibited increased leakage, especially below 40% (dry weight basis, DWB) seed moisture, the critical water content for viability loss in Spartina alterniflora. When isolated embryos were dried, the pattern and extent of increased solute leakage were similar in both Spartina species, even though S. pectinata embryos were >95% viable and S. alterniflora embryos were < 20% viable. We conclude that increased solute leakage is an artefact of embryo excision and not an accurate indicator of desiccation damage to recalcitrant Spartina alterniflora seeds.

Assessments of seed storage physiology among Arecaceae (palm) species are often inconclusive because seeds exhibit diverse responses to low temperature and moisture conditions. Interrelationships between dry matter accumulation, cell structure and water relations during seed development of the endangered Hawaiian endemic palm, Pritchardia remota, suggest that damage from drying results from mechanical strain. Endosperm and fruits accumulate dry mass through most of the 400 d gestation period, but embryos reached maximum dry mass about 250 d post-anthesis (DPA). Mostly sucrose and some triacylglycerols accumulated in the cytoplasm and vacuoles of embryo cells, and organelles in mature embryo cells de-differentiated. Water content and water potential decreased as embryos matured and embryos contained about 0.45 g H2O (g dry mass)− 1 ( − 26 MPa) at shedding. Mature embryos survived drying to 0.16 g g− 1 ( − 49 MPa), but further drying was lethal. A model of allowable cell shrinkage is consistent with the substantial, but incomplete, desiccation tolerance acquired in P. remota embryos, and provides a new framework to explain variation in critical water contents as embryos develop. We suggest that desiccation tolerance, which distinguishes recalcitrant and orthodox physiologies among seeds, can be quantified by mechanical strain when embryo cells shrink during drying.

Dehydrins are late embryogenesis abundant (LEA) proteins that accumulate during seed maturation and in response to abiotic stresses in vegetative tissues. They are thought to protect cellular components from dehydration stress. However, whether they play a role in survival in the dry state is not clear. In this study, an RNA interference (RNAi)-construct against the seed-expressed dehydrin of Arabidopsis thaliana, LEA14 (At2g21490), was introduced to wild-type plants, which led to a strong reduction in transcript abundance of the target gene as well as that of two other seed-expressed dehydrin homologues, XERO1 (At3g50980) and RAB18 (responsive to abscisic acid 18, At5g66400) in the transformants. Mature, dry seeds from the RNAi plants germinated to at least 95% after rehydration, indicating that seed desiccation tolerance was not affected, while they exhibited a twofold reduction in longevity. When stored at 75% relative humidity and 35°C, the seeds of two independent RNAi lines lost 50% of their viability in 10 d and 5 d, respectively, while it took 17 d for wild-type seeds to lose 50% viability. In addition, when seeds were imbibed in the presence of 100 mM NaCl, the seeds of RNAi plants exhibited reduced germination compared to wild-type seeds, suggesting that at least one of the three seed-specific dehydrins plays a role both against deterioration during storage at low moisture content and when imbibed tissues are submitted to salt stress at high moisture.

The potential longevity of japonica rice (Oryza sativa L. subsp. japonica) seed is particularly sensitive to high temperature – and thus climate change – during development and maturation. Cultivar Taipei 309 was grown at 28/20°C (12 h/12 h) and then from 19 DAA (days after 50% anthesis), when seeds were just over half filled, at 28/20°C, 30/22°C, 32/24°C or 34/26°C (12 h/12 h). Whereas ability to germinate ex planta had been achieved in almost all seeds by 24 DAA, only half the population were desiccation tolerant. Desiccation tolerance continued to increase over the subsequent 28 d, similarly at all four temperatures. Subsequent longevity, assessed by p50 (period in days to reduce viability to 50% in hermetic storage at 40°C with c. 15% moisture content), increased progressively at 28/20°C until 38 DAA, and remained constant until the final harvest (52 DAA). The three warmer temperature regimes provided similar longevity to 28/20°C at any one harvest, except at 38 DAA where the warmest (34/26°C) was poorer. That temperature regime also provided greater seed-to-seed variability within each survival curve. The results confirm that appreciable improvement in seed quality occurs during seed development and also subsequent maturation in japonica rice, but that increase in temperature from 28/20°C to 34/26°C during late seed filling onwards has comparatively little effect thereon. Comparison with previous investigations suggests that seed quality development may be less sensitive to high temperatures during late development and maturation than during the early seed development that precedes it.

The survival capacities of Muellerius capillaris (Nematoda: Protostrongylidae) free-living larval stages (L1) of an Israeli desert isolate (Nubian isolate) (92 mm rain per annum) and a French isolate from a temperate habitat (650 mm rain per annum) were compared under humid and extreme dry conditions. Under the dry conditions (7% relative humidity [r.h.] and 33% r.h. at 23 °C), both isolates exhibited the same remarkable survival capacity for periods of 17 and 28 days, respectively (>92% survival, P>0·1). However, during and after recovery from the anhydrobiotic state, different behaviour patterns of L1 were observed in each isolate. Under humid conditions (97% r.h.) for 10 days, both isolates exhibited similarly low survival percentages (13·4±4·6%, S.E., survival for the Nubian isolate, 3±3% for the French isolate, P>0·05). In water, the French isolate was less active, survived better (52±8·4% vs 28·7±3·7, by day 28, P<0·05) and retained a better morphological appearance (shrunk less) than the Nubian isolate. Larval development of the French isolate in the land snail Theba pisana was significantly faster than that of the Nubian isolate, reaching a higher proportion of infective stages (L3) on day 36 post-infection (78·6% vs 56% P>0·05). However, similar infection intensities were recovered in snails infected with either of the isolates (59·8±10·3, S.E., n=14, for the French isolate, 53±6, n=13, for the Nubian isolate, P<0·05).

This article reviews mechanisms by which specialized cells of different life forms have overcome the lethal effects of dehydration and considers how the maintenance of genetic information is central to survival. As a dynamic and hydrated molecule in vivo, DNA can assume different conformational structures depending upon the water activity, the base sequence and the presence of specific binding proteins. The attainment of stable secondary structures that are resistant to degradation in vivo at low water potentials is proposed as a likely accompaniment to desiccation tolerance. In addition, chemical modification of bases in DNA, the extent of methylation and conformational changes could determine the expression of different gene sequences as cells pass from desiccation-tolerant to desiccation-intolerant states. We monitored the integrity of extracted DNA in embryos of seeds and in wind-dispersed pollen during transition from their desiccation tolerance to desiccation intolerance on hydration and germination. We present evidence to show that the DNA of these two stages is different and that it is the DNA from desiccation-tolerant cells only that retains integrity when the cells are subjected to desiccation regimes. We discuss these findings in relation to certain hydration-sensitive DNA structures and to other relevant biological systems.

In view of the finding that dry seeds may exist in the glassy state, a brief review of the commonest methods of detection and quantification of the glass transition is presented. While the glassy state may contribute to seed tolerance of desiccation, it does not appear to account for desiccation tolerance. Its major function in dry seeds may be its contribution to the stability of the seed components during storage.

The fruits of Quercus robur are shed containing seeds at high moisture contents which remain desiccation sensitive and exhibit recalcitrant storage behaviour. Little is known of the control of seed development in these and other recalcitrant seeds. In the present work the changing concentrations of four plant growth regulators during seed reserve accumulation was studied over five years on seeds from the same tree. The pattern of reserve accumulation and changing ABA content in seeds differed between years. Although ABA content in the cotyledons increased in line with increasing dry weight to different contents at shedding, similar concentrations existed in cotyledons in each year. Thus ABA did not appear to be influencing dry weight accumulation. However, unlike orthodox seeds the decline in ABA concentration prior to shedding was limited and consistent with a continuing role for ABA in preventing precocious germination. An earlier peak in ABA concentration was associated with greater desiccation tolerance at shedding across years. The concentrations of zeatin and zeatin riboside in cotyledons were similar in each year and declined during reserve accumulation in a similar fashion to that reported for orthodox seeds. By contrast, IAA concentration increased in both the cotyledons and axes in the latter stages of seed development, opposite to that reported for orthodox seeds. It is possible that the increasing IAA concentration in cotyledons and axes and the stable concentration of zeatin and zeatin riboside throughout the latter stages of development in the axes of Q. robur are linked to the maintenance of active metabolism for the rapid initiation of germination upon shedding observed in seeds of this species.

In the presence of abscisic acid (ABA), contents of glucose and fructose decreased in carrot embryoids, whereas umbelliferose increased during dehydration. The acquisition of desiccation tolerance was tested by scoring germinated embryoids. It did not keep pace with the changes in soluble carbohydrate contents during development, which suggests that in these somatic embryoids di- and oligosaccharides are not the determining factors for anhydrobiosis. However, on slow dehydration, umbelliferose increased, while sucrose decreased and the monosaccharides declined completely. These changes were positively correlated with increased desiccation tolerance. A similar analysis of slowly dried, ABA-treated alfalfa embryoids showed that stachyose and sucrose increased and the monosaccharides declined with the acquisition of desiccation tolerance. These data support the contention that carbohydrates are involved in anhydrobiosis.

Loss of vigour in leek (Allium porrum L.) seed lots is accompanied by the appearance of damage to ribosomal RNA in quiescent embryo tissue. Polyethylene glycol osmopriming treatments of such low-vigour seed permit replacement of this damaged ribosomal rRNA over a 7-day priming period. Low-vigour leek seeds germinated for 6 days without a prior osmopriming treatment still exhibit evidence of damaged ribosomal RNA in embryo tissue. Osmoprimed leek seeds dried back whilst still in the desiccation-tolerant state retain most of the benefits conferred by priming treatments. Osmoprimed leek seeds dried back when having reached the desiccation-sensitive state germinate poorly, if at all, and exhibit much reduced rates of protein synthesis in embryo tissue upon rehydration compared to desiccation-tolerant seeds. Embryo tissue from dried back, desiccation-tolerant seeds exhibits the capacity for much higher levels of protein synthesis than embryo tissue from unprimed seeds at equivalent stages of imbibition. RNA levels continue to increase in embryo tissue upon rehydration of dried back, desiccation-tolerant leek seeds but not in desiccation-sensitive seeds. Loss of protein synthetic capacity in desiccation-sensitive seeds during rehydration is accompanied by increasing levels of ribosomal RNA degradation. Such ribosomal RNA degradation may be indicative of the germinative capacity of leek seeds dried back after osmopriming treatments.

The interactions between water content and temperature on freezing and desiccation damage were examined for Zizania (wild rice) embryos at several stages of maturity. The water content of excised embryos was manipulated by flash drying at 35°C or room temperature to between 2.5 g H2O/g dw and 0.05 g/g. Embryos were then exposed to temperatures ranging from 5 to −50°C. Viability following the drying and cooling treatments was assayed by leakage of electrolytes and germination in culture. Viability of embryos decreased when embryos were dried below a critical water content. The critical water content was greatest for the least mature embryos. Critical water contents were also temperature dependent and increased with decreasing temperature. Even though the critical water content varied with developmental status and temperature, the water activity corresponding to the critical water content appeared to be constant at 0.90. The most mature embryos survived temperatures as low as −50°C while the least mature embryos survived only to −18°C. These trends were predicted by ‘phase diagrams’ based on the physical properties of water in embryos at different stages of maturity (Vertucci et al., 1994a). Our results confirm the earlier prediction that long term preservation of Zizania grains is possible at −20°C and the degree of success will be related to the maturity status of the embryos.

It is proposed that desiccation tolerance in the embryo of seeds depends upon the capacity to repair damage to genomic DNA when the desiccated embryo is rehydrated. From a study of imbibed and hydrated embryos of rye (Secale cereale) and wild oat (Avena fatua) evidence is provided that it is neither the extent of water uptake by the cells, the ensuing stability of the DNA to desiccation, nor the onset of S-phase DNA synthesis in the first cell cycle of germination that determines whether the desiccated embryo will survive. It is shown that when α- and β-polymerases of DNA repair are inhibited by aphidicolin and dideoxythymidine-5'-triphosphate, respectively, a γ-irradiation-induced DNA fragmentation cannot be fully repaired. It is shown that in hydrated embryos, at a stage when desiccation tolerance is lost, embryo cells still repair irradiation-induced damage, but this repaired DNA is unstable to desiccation and cannot be rerepaired when water is again made available. The failure to re-repair on rehydration appears to be critical to embryo survival and successful germination.

Norway maple (Acer platanoides L.) seeds were harvested at different stages of seed development and maturation in 1989–91. As maturation drying progressed, the seed populations showed increasing desiccation tolerance: at 67–69% moisture content, no seeds survived desiccation below 10% moisture content; maturation drying to 55–57% moisture content (values corresponding with the end of the seed-filling phase) improved desiccation tolerance, but nevertheless most seeds were unable to withstand desiccation to 5–7% moisture content; further maturation drying to 27–28% moisture content enabled the seeds to survive considerable desiccation, no loss in viability occurring in seeds dried to 3% moisture content. This considerable improvement in desiccation tolerance after the end of the seed-filling phase was correlated (P<0.05) with the progress of maturation drying and may be associated with the increase in the potential longevity of seeds of other species that occurs during seed development subsequent to seed filling.

Excised Helianthus annuus embryos were capable of precocious germination as early as 7 d after pollination. At this stage they were not capable of withstanding desiccation treatment. The desiccation tolerance was acquired at a stage of development which depended on the rate of water loss to which embryos were subjected. Coincident with the acquisition of tolerance to a rapid desiccation, they gained the capacity to withstand 24 h preincubation in agitated water; this acquisition could be correlated to a sharp decrease of the leaching of electrolytes, amino acids and proteins in the incubation water, suggesting that a change in membrane properties played a key role in this transition. Moreover, the acquisition of tolerance to these stresses was coincident with the increase of endogenous abscisic acid (ABA). ABA applied in vitro to young embryos induced tolerance both to desiccation and to preincubation in water: this in vitro acquisition of tolerance was again correlated with a decrease in the leakage.

The desiccation tolerance of Zizania palustris seeds has been debatable, but recently it has emerged that survival to low moisture contents is a function of the temperature during dehydration. Survival of dehydration is greatest at 25–30°C and viability declines as the dehydration temperature is reduced. On the other hand, extended hydrated chilling (stratification) is required to break dormancy. The present contribution examines the ultrastructural condition of embryonic axes of Zizania palustris after dehydration at various temperatures, and following reimbibition and stratification. Axis cells sustained least damage when dehydration was carried out at 25°C and ultrastructural deterioration was more severe with lower temperatures during water loss. Damage sustained as a result of unfavourably low dehydration temperatures was visible when seeds were fixed from the dry state and was generally exacerbated during fully imbibed stratification. However, 25°C represented the optimum dehydration temperature; seeds that had been dried at 30°C also showed considerable ultrastructural disturbance when fixed from the dry state. This was largely reversed during fully imbibed stratification, although signs of the damage that had been sustained still persisted. These observations are in keeping with the germination behaviour of Z. palustris seeds dehydrated at temperatures above and below 25°C. The results are discussed in terms of the predominance of one of two mechanisms of membrane deterioration, depending on the dehydration temperature, which are probably not mutually exclusive. It is hypothesized that, at temperatures above 25°C, damage by free-radical-mediated events may predominate, whereas, at temperatures below the optimum, irreversible lipid-phase transitions may be the major factor resulting in membrane damage.