From the end to the start : genetic and physiological aspects of the lifespan of seeds

Seed dormancy and longevity contribute to seed quality and together define the lifespan of the seed. Seeds are often stored dry in research and agriculture and during this dry storage, dormancy is released and ultimately the seeds deteriorate (up to 10 years for Arabidopsis). The mechanisms underlying seed lifespan and the interaction between dormancy and longevity are not fully understood.       First, a new method (elevated partial pressure of oxygen, EPPO) was developed to be able to mimic and accelerate the entire seed lifespan. With this method, quantitative trait loci (QTL) mapping could be performed and candidate genes underlying two longevity QTL have been identified. A third longevity locus was identified by genome wide association studies, and knock-out lines for the candidate genes were analysed with the EPPO method.       Dormancy was also studied in a field experiment. Transcriptome analysis was performed on seeds at different stages during dormancy cycling to gain more insight into the molecular mechanisms. The transitions between dormancy levels during cycling were found to be accompanied by large changes in the transcriptome. The genes encoding for the translation machinery are upregulated in non-dormant seeds in the soil, indicating that these seeds are prepared for rapid germination. &nbsp

A Perspective on Secondary Seed Dormancy in Arabidopsis thaliana

Primary seed dormancy is the phenomenon whereby seeds newly shed by the mother plant are unable to germinate under otherwise favorable conditions for germination. Primary dormancy is released during dry seed storage (after-ripening), and the seeds acquire the capacity to germinate upon imbibition under favorable conditions, i.e., they become non-dormant. Primary dormancy can also be released from the seed by various treatments, for example, by cold imbibition (stratification). Non-dormant seeds can temporarily block their germination if exposed to unfavorable conditions upon seed imbibition until favorable conditions are available. Nevertheless, prolonged unfavorable conditions will re-induce dormancy, i.e., germination will be blocked upon exposure to favorable conditions. This phenomenon is referred to as secondary dormancy. Relative to primary dormancy, the mechanisms underlying secondary dormancy remain understudied in Arabidopsis thaliana and largely unknown. This is partly due to the experimental difficulty in observing secondary dormancy in the laboratory and the absence of established experimental protocols. Here, an overview is provided of the current knowledge on secondary dormancy focusing on A. thaliana, and a working model describing secondary dormancy is proposed, focusing on the interaction of primary and secondary dormancy

Evolutionary ecophysiology of seed desiccation sensitivity

Desiccation sensitive (DS) seeds do not survive dry storage due to their lack of desiccation tolerance. Almost half of the plant species in tropical rainforests produce DS seeds and therefore the desiccation sensitivity of these seeds represents a problem for and long-term biodiversity conservation. This phenomenon raises questions as to how, where and why DS (desiccation sensitive)-seeded species appeared during evolution. These species evolved probably independently from desiccation tolerant (DT) seeded ancestors. They adapted to environments where the conditions are conducive to immediate germination after shedding, e.g. constant and abundant rainy seasons. These very predictable conditions offered a relaxed selection for desiccation tolerance that eventually got lost in DS seeds. These species are highly dependent on their environment to survive and they are seriously threatened by deforestation and climate change. Understanding of the ecology, evolution and molecular mechanisms associated with seed desiccation tolerance can shed light on the resilience of DS-seeded species and guide conservation efforts. In this review, we survey the available literature for ecological and physiological aspects of DS-seeded species and combine it with recent knowledge obtained from DT model species. This enables us to generate hypotheses concerning the evolution of DS-seeded species and their associated genetic alterations

Evaluating the EPPO method for seed longevity analyses in Arabidopsis

Seed longevity (storability) is an important seed quality trait. High seed quality is important in agriculture, for the industry, and for safeguarding biodiversity as many species are stored as seeds in genebanks. To ensure ex-situ seed survival, seeds are mostly stored at low relative humidity and low temperature. Oxidation is the main cause of seed deterioration in these dry storage conditions. The molecular mechanisms underlying dry seed survival remain poorly understood. Research on seed longevity is hampered by the lack of an experimental ageing method that mimics dry ageing well. Here, we propose the Elevated Partial Pressure of Oxygen (EPPO) method as the best available method to mimic and accelerate dry seed ageing. We have tested seed germination in Arabidopsis thaliana after EPPO storage at two different relative humidity (RH) conditions and confirm the large effect of oxygen and the seed moisture content on ageing during dry storage. Comparative Quantitative trait locus (QTL) analysis shows that EPPO at 55 % RH mimics dry ageing better than the commonly used Artificial Ageing and Controlled Deterioration tests at higher moisture levels.</p

Altitudinal and climatic associations of seed dormancy and flowering traits evidence adaptation of annual life cycle timing in Arabidopsis thaliana

The temporal control or timing of the life cycle of annual plants is presumed to provide adaptive strategies to escape harsh environments for survival and reproduction. This is mainly determined by the timing of germination, which is controlled by the level of seed dormancy, and of flowering initiation. However, the environmental factors driving the evolution of plant life cycles remain largely unknown. To address this question we have analysed nine quantitative life history traits, in a native regional collection of 300 wild accessions of Arabidopsis thaliana. Seed dormancy and flowering time were negatively correlated, indicating that these traits have coevolved. In addition, environmental-phenotypic analyses detected strong altitudinal and climatic clines for most life history traits. Overall, accessions showing life cycles with early flowering, small seeds, high seed dormancy and slow germination rate were associated with locations exposed to high temperature, low summer precipitation and high radiation. Furthermore, we analysed the expression level of the positive regulator of seed dormancy DELAY OF GERMINATION 1 (DOG1), finding similar but weaker altitudinal and climatic patterns than seed dormancy. Therefore, DOG1 regulatory mutations are likely to provide a quantitative molecular mechanism for the adaptation of A. thaliana life cycle to altitude and climate.</p

SIEVE ELEMENT-LINING CHAPERONE1 restricts aphid feeding on arabidopsis during heat stress

The role of phloem proteins in plant resistance to aphids is still largely elusive. By genome-wide association mapping of aphid behavior on 350 natural Arabidopsis thaliana accessions, we identified the small heat shock-like SIEVE ELEMENT-LINING CHAPERONE1 (SLI1). Detailed behavioral studies on near-isogenic and knockout lines showed that SLI1 impairs phloem feeding. Depending on the haplotype, aphids displayed a different duration of salivation in the phloem. On sli1 mutants, aphids prolonged their feeding sessions and ingested phloem at a higher rate than on wild-type plants. The largest phenotypic effects were observed at 26°C, when SLI1 expression is upregulated. At this moderately high temperature, sli1 mutants suffered from retarded elongation of the inflorescence and impaired silique development. Fluorescent reporter fusions showed that SLI1 is confined to the margins of sieve elements where it lines the parietal layer and colocalizes in spherical bodies around mitochondria. This localization pattern is reminiscent of the clamp-like structures observed in previous ultrastructural studies of the phloem and shows that the parietal phloem layer plays an important role in plant resistance to aphids and heat stress

NADP-MALIC ENZYME 1 Affects Germination after Seed Storage in Arabidopsis thaliana

Aging decreases the quality of seeds and results in agricultural and economic losses. The damage that occurs at the biochemical level can alter the seed physiological status. Although loss of viability has been investigated frequently, little information exists on the molecular and biochemical factors involved in seed deterioration and loss of viability. Oxidative stress has been implicated as a major contributor to seed deterioration, and several pathways are involved in protection against this. In this study, we show that seeds of Arabidopsis thaliana lacking a functional NADP-MALIC ENZYME 1 (NADP-ME1) have reduced seed viability relative to the wild type. Seeds of the NADP-ME1 loss-of-function mutant display higher levels of protein carbonylation than those of the wild type. NADP-ME1 catalyzes the oxidative decarboxylation of malate to pyruvate with the simultaneous production of CO2 and NADPH. Upon seed imbibition, malate and amino acids accumulate in embryos of aged seeds of the NADP-ME1 loss-of-function mutant compared with those of the wild type. NADP-ME1 expression is increased in imbibed aged as compared with non-aged seeds. NADP-ME1 activity at testa rupture promotes normal germination of aged seeds. In seedlings of aged seeds, NADP-ME1 is specifically active in the root meristematic zone. We propose that NADP-ME1 activity is required for protecting seeds against oxidation during seed dry storage