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Seeds offer a unique perspective from which to view biology. An individual seed is an autonomous biological entity that must rely on its own resources (and resourcefulness) to persist after dispersal and to time its transition to germination and seedling growth to coincide with environmental opportunities for survival. At the same time, seed biology in agriculture and ecology is determined largely by the behaviours of populations of individual seeds. The percentage of seeds in a population that is in a particular state (e.g. dormant, germinated, dead) at a given time is a fundamental metric of seed biology. This duality of individual diversity underlying consistent population-wide behaviour patterns can be described quantitatively using population-based threshold (PBT) models. While conceptually simple, these models are highly flexible and can describe the wide diversity of responses of seed populations to temperature, water potential, hormones, oxygen, light, ageing and combinations of these factors. This seed behaviour is linked to respiratory rates of individual seeds, indicating that basic metabolic processes within seeds vary among individuals in accordance with PBT principles. Looking more broadly across microbial, plant and animal biology, examples of cellular diversity in hormonal sensitivity, gene expression, developmental responses and signalling abound. This variation often is termed ‘noise’, and analysis efforts are focused on extracting mean signals from this variation to understand regulatory pathways. However, extension of the PBT approach to the cellular and molecular levels suggests that population sensitivity distributions and recruitment phenomena may underlie many fundamental biological processes. Thus, concepts and quantitative approaches developed for the analysis of seed populations can be applied across biological scales from molecules to ecosystems to interpret inherent biological variation and provide mechanistic insights into the nature of biological regulatory systems.
Knowledge and prediction of seasonal weed seedling emergence patterns is useful in weed management programs. Seed dormancy is a major factor influencing the timing of seedling emergence, and once dormancy is broken, environmental conditions determine the rate of germination and seedling emergence. Seed dormancy is a population-based phenomenon, because individual seeds are independently sensing their environment and responding physiologically to the signals they perceive. Mathematical models based on characterizing the variation that occurs in germination times among individual seeds in a population can describe and quantify environmental and after-ripening effects on seed dormancy. In particular, the hydrothermal time model can describe and quantify the effects of temperature and water potential on seed germination. This model states that the time to germination of a given seed fraction is inversely proportional to the amount by which a given germination factor (e.g., temperature or water potential) exceeds a threshold level for that factor. The hydrothermal time model provides a robust method for understanding how environmental factors interact to result in the germination phenotype (i.e., germination pattern over time) of a seed population. In addition, other factors that influence seed dormancy and germination act by causing the water potential thresholds of the seed population to shift to higher or lower values. This relatively simple model can describe and quantify the germination behavior of seeds across a wide array of environmental conditions and dormancy states, and can be used as an input to more general models of seed germination and seedling emergence in the field.
Seed germination is responsive to diverse environmental, hormonal and chemical signals. Germination rates (i.e. speed and distribution in time) reveal information about timing, uniformity and extent of germination in seed populations and are sensitive indicators of seed vigour and stress tolerance. Population-based threshold (PBT) models have been applied to describe germination responses to temperature, water potential, hormones, ageing and oxygen. However, obtaining detailed data on germination rates of seed populations requires repeated observations at frequent times to construct germination time courses, which is labour intensive and often impractical. Recently, instruments have been developed to measure repeatedly the respiration (oxygen consumption) of individual seeds following imbibition, providing complete respiratory time courses for populations of individual seeds in an automated manner. In this study, we demonstrate a new approach that enables the use of single-seed respiratory data, rather than germination data, to characterize the responses of seed populations to diverse conditions. We applied PBT models to single-seed respiratory data and compared the results to similar analyses of germination time courses. We found consistent and quantitatively comparable relationships between seed respiratory and germination patterns in response to temperature, water potential, abscisic acid, gibberellin, respiratory inhibitors, ageing and priming. This close correspondence between seed respiration and germination time courses enables the use of semi-automated respiratory measurements to assess seed vigour and quality parameters. It also raises intriguing questions about the fundamental relationship between the respiratory capacities of seeds and the rates at which they proceed toward completion of germination.
Lettuce (Lactuca sativa L.) and onion (Allium cepa L.) seeds have relatively short longevity during storage and their germination is sensitive to environmental stress. Seed priming (controlled hydration followed by drying) can improve seed germination under stressful conditions, inducing faster and more uniform germination over broader temperature ranges, but it can also reduce seed longevity in storage. Controlled deterioration (CD) tests are often employed to study longevity by ageing seeds rapidly at elevated temperature and moisture content, and primed seeds are particularly sensitive to CD conditions. As reactive oxygen (O2) species are thought to be involved in seed deterioration, we tested whether storage under reduced O2 atmospheres (0 and 2% O2) would extend the longevity of primed and non-primed seeds under low relative humidity (RH) (33% RH+37°C) and CD (75% RH+50°C) storage conditions. The longevity of both non-primed and primed lettuce seeds in low RH storage was extended by anaerobic environments, but the effect of O2 was much less under CD conditions. In onion, only primed seeds exhibited a beneficial effect of low O2 atmospheres under both types of ageing conditions. In both species, storage under anaerobic conditions was beneficial for extending the longevity of primed seeds, but was not able to ameliorate fully the negative effect of priming on storage life.
A recent hypothesis states that the accumulation of amino-carbonyl reaction products of reducing sugars with proteins (Maillard products) is related to the loss of seed vigour and viability during ageing. Since Maillard products are biochemical end products, their accumulation should provide an index of seed deterioration. A simple fluorescence assay has been used to estimate the quantity of Maillard products in aqueous seed extracts. We tested whether an accumulation of fluorescent compounds correlates with the loss of seed viability during ageing. Fluorescence of extracts from lettuce (Lactuca sativa) embryonic axes increased after controlled deterioration, but was not correlated with ageing period. Fluorescence of extracts from whole broccoli (Brassica oleracea var. italica) seeds was either unchanged or declined during controlled deterioration. Extracts from naturally aged whole seeds of carrot (Daucus carota), tomato (Lycopersicon esculentum), and cauliflower (B. oleracea var. botrytis) had higher fluorescence than controls, but there was no consistent increase in fluorescence in proportion to the loss of viability. Extracts of naturally aged onion (Allium cepa) seeds exhibited less fluorescence than extracts from seeds with higher viability. We conclude that the fluorescence assay does not have general utility as an index of seed deterioration.
Lettuce (Lactuca sativa L.) seeds are inhibited from germinating above an upper temperature limit that is dependent upon cultivar, growing conditions and seed treatments. Thermoinhibition is accompanied by an increasing sensitivity of germination to reduced water potentials (ψ). We have employed a water relations analysis (hydrotime model) of seed germination rates to investigate the basis of thermoinhibition. Germination rates can be characterized by the distribution of base water potentials among seeds in the population (ψb(g)) and a hydrotime constant indicating the total accumulated hydrotime (MPa · h) above the base required for radicle emergence. The hydrotime model adequately described germination time courses across a range of ψ at both high and low temperatures. Increasing temperature caused the ψb(g) distributions to become more positive, accounting for the greater sensitivity to ψ and inhibition of germination. Increases in embryo osmotic potential and in the turgor yield thresholds of both the radicle and the endosperm/pericarp envelope contributed to this change. Seed priming (prehydration and drying) treatments speeded germination by reducing the hydrotime requirement. Priming also resulted in smaller increases in ψb(g) at high temperature, alleviating thermoinhibition by lowering the embryo yield threshold sufficiently to compensate for the increased endosperm resistance. The beneficial effects of priming in lettuce appear to occur primarily in the embryo, rather than in the surrounding envelope tissues.
Dehydrin and QP47, proteins present in mature pea seeds (Pisum sativum), have been proposed to play protective roles during desiccation. To identify possible relationships between these proteins and desiccation tolerance, their tissue locations and patterns of synthesis and degradation have been examined during germination. Tissue locations were determined by immunocytochemistry using polyclonal antibodies raised against a conserved dehydrin amino acid sequence and against purified QP47. In embryonic axis and cotyledon cells, QP47 and dehydrin were distributed uniformly with no apparent nuclear or organellar specificity. Both proteins were present in 24 h-imbibed axes that had not initiated radicle growth but were completely absent from 24 h-imbibed axes that had begun to grow. The amounts of QP47 and dehydrin in embryonic axes decreased with time after the start of imbibition and were undetectable by 48 h. When germination was prevented by polyethylene glycol (PEG) or abscisic acid (ABA), both proteins remained at their original amounts. Thus, both QP47 and dehydrin disappeared coincidently with the beginning of growth and not simply as a function of the time after imbibition. QP47 persisted in cotyledons until at least 31 days into seedling growth, whereas dehydrin was not detectable in cotyledons after 7 days. Dehydrin, but not QP47, could be re-induced in pea shoots and cotyledons by dehydration. The timing of degradation of both proteins was correlated with the loss of desiccation tolerance during germination of pea axes.
The papers in this special section of Seed Science Research are products of a symposium on Seed Biology and Technology: Applications and Advances, held in Fort Collins, Colorado, on 13–16 August, 1997. The symposium was convened as a cooperative effort of Regional Research Project W-168 within the United States Department of Agriculture (USDA) Cooperative States Research, Extension and Education Service (CSREES) system. Regional Research Projects are authorized by the Hatch Act, which established the Agricultural Experiment Station (AES) system in the United States (US Code). This is a system in which land-grant institutions in each state conduct research and education programmes relevant to agriculture, the environment and society. Regional Research projects are a mechanism ‘for cooperative research in which two or more State agricultural experiment stations are cooperating to solve problems that concern the agriculture of more than one state.’ Such projects ‘can provide the solution to a problem of fundamental importance or fill an important gap in our knowledge from the standpoint of the present and future agriculture of the region’
A water relations (hydrotime) model was employed to characterize the responses of lettuce (Lactuca sativa L.) seed germination to temperature, water potential (ψ), endosperm disruption, and the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC). The base water potentials (ψb), standard deviations of base water potential and hydrotime constants (θH) estimated by the model were used to predict the germination rates at reduced ψ and supraoptimal temperatures in the presence or absence of ACC. The distribution of ψb values among seeds in the population (ψb(g)) increased (became more positive) as the imbibition temperature increased, indicating a greater sensitivity of seeds to reduced ψ at thermoinhibitory temperatures. Slitting the seeds at the cotyledonary end to disrupt the integrity of the endosperm extended the high temperature range for germination and maintained lower ψb(g) distributions compared with intact seeds as temperature increased. ACC (10 mm) in the imbibition solution raised the upper temperature limit for germination by about 2°C, and also lowered ψb(g) values by 0.1 to 0.2 MPa. The effect of ACC on ψb(g) was more pronounced at higher imbibition temperatures and in slit seeds. ACC (via conversion to ethylene) extends the high temperature limit for lettuce seed germination by acting in the embryo to maintain a lower ψ threshold for the initiation of growth as temperature increases. Evidence from other species as well suggests that this may be a general mechanism for the action of ethylene in promoting seed germination.
Seed germination rates are sensitive to both temperature (T) and water potential (ψ). The times to germination of seeds imbibed at suboptimal T and/or reduced ψ are inversely proportional to the amounts by which T exceeds a base temperature (Tb) and ψ exceeds a base water potential (ψb). Germination rates across a range of suboptimal T and ψ can be normalized on the basis of the hydrothermal time accumulated in excess of these thresholds. However, seeds can also progress metabolically toward germination even at T or ψ too low to allow radicle emergence to occur. Seeds preimbibed at low ψ and dried back, or primed, germinate more rapidly upon subsequent reimbibition. We show here that the increase in germination rates of tomato (Lycopersicon esculentum Mill.) seeds resulting from seed priming is linearly related to the hydrothermal time accumulated during the priming treatment. The threshold temperature (Tmin = 7.05°C) and water potential (ψmin = −2.50 MPa) for metabolic advancement were considerably lower than the corresponding thresholds for radicle emergence of the same seed lot (Tb = 11°C; ψb = −0.71 MPa), allowing the accumulation of hydrothermal priming time that is subsequently expressed as more rapid germination when T or ψ increase. The hydrothermal time model can now be applied to quantify and analyse germination rates of seeds across the entire range of suboptimal T and ψ at which metabolic progress toward radicle emergence is possible.
Both temperature (T) and water potential (ψ) have consistent and quantifiable effects on the rate and extent of seed germination (radicle emergence). Germination at suboptimal T can be characterized on the basis of thermal time, or the T in excess of a base (Tb) multiplied by the time to a given percentage germination (tg). Similarly, germination at reduced ψ can be characterized on a hydrotime basis, or the ψ in excess of a base (ψb) multiplied by tg. Within a seed population, the variation in thermal times to germination for a specific percentage (g) is based upon the normal distribution of ψb values among seeds (ψb(g)). Germination responses across a range of suboptimal T and ψ might be accounted for by a general hydrothermal time model incorporating both T and ψ components. We tested this hypothesis for tomato (Lycopersicon esculentum Mill.) seeds of two genotypes differing in germination rates and tolerance of suboptimal T and ψ. For combinations of T (10−25°C) and ψ (0 to −0.9 MPa), a general hydrothermal time model accounted for approximately 75% of the variation in times to germination within the seed populations of both genotypes, and over 96% of the variation in median germination rates. However, ψb(g) distributions were sensitive to both the T and ψ of imbibition, resulting in a poor fit of the model to specific time course data. Analysis of germination timing separately for low and high ψ ranges within a given T resulted in specific models accounting for 88−99% of the variation in individual germination times and >99% of the variation in madian germination rates. Thus, for a given T and ψ range, the hydrotime model closely matched tomato seed germination time courses. Accumulated hydrothermal time accounted well for germination rates at ψ> −0.5 MPa across suboptimal T if ψb(g) was allowed to vary with T. Germination did not show a consistent response to T at ψ < −0.5 MPa, and estimated Tb values varied over different T ranges. Generalization of the hydrothermal time model across the entire range of suboptimal T and ψ was limited by physiological adjustments of the seeds to their current environment. The hydrothermal time model detected and quantified these adjustment processes that would otherwise not be evident from inspection of germination time courses. Temperature and water potential influence the time to germination via physiological mechanisms that reciprocally interact.
Seeds vary widely in the sensitivity of germination to oxygen (O2) partial pressure, depending upon the species, temperature, dormancy state and physiological status of the seeds. Most analyses of the O2 sensitivity of germination have focused on final germination percentages and estimated the O2 percentage in air that is required to reduce germination to a given percentage (usually 50%). In contrast, we have applied a population-based threshold model utilizing time courses of germination to quantify three parameters related to seed germination sensitivity to O2 availability: the median base (or threshold) O2 percentage, the standard deviation of O2 thresholds among seeds in the population, and an oxygen–time constant that relates O2 percentage to germination timing. The model fits germination responses accurately across a wide range of O2 concentrations. The response to O2 was logarithmic in all cases, with the O2 percentage required for 50% germination ranging from 21% to as low as 0.005%, depending upon the species, the temperature and the seed dormancy level. Modelling indicated that some seeds can adapt to low O2 percentages and shift their thresholds to lower values over time. Lower temperatures decreased the minimum O2 threshold, as did after-ripening. Seed priming generally reduced the oxygen–time constant and increased the standard deviation of germination responses, but had relatively little effect on the O2 sensitivity per se. The population-based threshold model can be used to quantify the O2 sensitivity of seed germination and to predict germination rates and percentages when O2 availability is limiting.
Metabolic heat and CO2 production rates were measured by microcalorimetry during germination of two melon (Cucumis melo L.) accessions, Noy Yizre’el (a cold-sensitive cultivar) and Persia 202 (a cold-tolerant breeding line), differing in their ability to germinate at low temperature (14°C). In seeds that were imbibed at either 14 or 25°C, both heat production rates (φ) and CO2 evolution rates (RCO2) WERE HIGHER IN PERSIA 202 COMPARED TO NOY YIZRE’EL. AT 14°C, Φ AND RCO2of intact Noy Yizre’el seeds remained close to zero and germination was inhibited, but metabolic activity increased with time in decoated seeds (testa removed) and most embryos germinated. The presence of the testa had less effect on metabolic activity or germination of Persia 202 seeds at 14°C. The ratio of φ/RCO2(calorespirometric ratio) is an indication of the substrate being utilized for respiration, with lower values (∼455 kJ mol–1) associated with carbohydrate substrates and higher values (∼800 kJ mol–1) associated with lipid substrates. Removal of the testae of Noy Yizre’el seeds increased φ/RCO2 at 14°C, suggesting that improved oxygen supply to the embryo promotes a shift from carbohydrate to lipid respiratory substrates associated with germination. Consistent with this interpretation, when the hilum apertures of the seed coats were sealed with silicone, φ, RCO2 and φ/RCO2of Noy Yizre’el seeds remained low and germination was inhibited at 25°C, while in Persia 202 seeds the same treatment had relatively little effect on φand RCO2, φ/RCO2approached the value expected for lipid respiratory substrates (700 kJ mol–1), and germination occurred. This confirms earlier anatomical work indicating that the testae of Persia 202 seeds were more porous than were those of Noy Yizre’el seeds, contributing to the greater tolerance of Persia 202 seeds to low temperatures. Abscisic acid (ABA) was also inhibitory to melon seed germination; imbibition of seeds at 25°C in 250 μM ABA decreased φ and RCO2, and φ/RCO2 remained lower than in control seeds in both genotypes. This effect of ABA on metabolic activity could be a cause or consequence of inhibition of germination.
Tomato seeds that have been dried, imbibed and redried (primed) develop internal free space between the embryo and endosperm. Seeds of the ABA-deficient sitiens(sitw) tomato mutant can exhibit internal free space at the completion of seed development even without priming. Both primed and sitwseeds germinate more rapidly than untreated wild-type seeds. To determine whether internal anatomy predicts germination physiology, individual sitwand primed wild-type seeds were sorted into three categories based upon the extent of internal free space observed non-destructively using X-radiography. Category 3 (C3, extensive free space present) sitw seeds completed germination more rapidly than all other seed categories and genotypes in water, in abscisic acid (ABA) or under far-red illumination. The force necessary to puncture the endosperm caps (and testa) of C3 sitw seeds was less, and the percentage of nuclei in C3 sitw radicle tips in the G2 stage of the cell cycle was greater than for all other seed categories. Wild-type seeds exhibited free space following long-term priming, but germination was still prevented by far-red light and ABA, and endosperm cap strength and nuclear DNA contents were not altered. Endo-β-mannanase activity of individual endosperm caps was not consistently related to their resistance to puncture. While internal free space is diagnostic for primed tomato seeds and occurs in a fraction of sitw seeds, it is not predictive of many aspects of germination physiology.
As seed dormancy is released within a seed population, both the rate and percentage of germination increase progressively with increasing dose of a dormancy-breaking treatment or condition. Population-based models can account for this behaviour on the basis of shifting response thresholds as dormancy is alleviated. In particular, hydrothermal time analysis of germination sensitivity to water potential (Ψ) and temperature (T) can describe these features of seed behaviour. We used the hydrothermal time model to analyse the effects of dormancy-breaking treatments on germination of dormant true (botanical) potato (Solanum tuberosum L.) seeds (TPS). After-ripening (37°C and 4% seed moisture content) of TPS for 7 or 30 days partially or fully alleviated primary dormancy. The median base water potential required to prevent germination [Ψb(50)] decreased from –0.25 MPa in control seeds to –0.87 MPa and –1.83 MPa after 7 and 30 days of after-ripening, respectively. In contrast, the base temperature for germination (Tb) was relatively unaffected (0–3.3°C). Fluridone (50 μM), an inhibitor of abscisic acid (ABA) biosynthesis, also promoted germination of dormant TPS and lowered Ψb(50), indicating a role for de novo synthesis of ABA during dormancy maintenance. Moist chilling (3 days at 4°C) or gibberellin (100 μM) alleviated secondary dormancy and lowered Ψb(50) values from –0.08 MPa to –0.36 and –0.87 MPa, respectively. The hydrothermal time model allows quantification of dormancy levels and explains why changes in germination speed and percentage are closely correlated during dormancy alleviation.
Correlative evidence indicates that sucrose and α-galactosyl-sucrose oligosaccharides (raffinose family oligosaccharides; RFOs) may be involved in seed longevity. Priming treatments (hydration in water or osmotic solutions followed by drying) can improve short-term seed performance but often result in reduced seed longevity. As RFOs are metabolized quickly following seed imbibition, loss of RFOs during priming could lead to more rapid deterioration in dry storage. This hypothesis was tested by measuring sucrose and oligosaccharide contents and potential longevity of primed seeds. Raffinose contents of whole lettuce (Lactuca sativa L.) seeds declined during hydration and priming and were correlated with decreased median potential viability (p50). However, this relationship was less significant when only the embryonic axes were analysed. In tomato (Lycopersicon esculentum Mill.) and impatiens (Impatiens balsamina L.) seeds, planteose was the major galactosyl-sucrose oligosaccharide and only small quantities of RFOs were present. Planteose contents declined during priming in seeds of both species, while sucrose contents increased or remained constant. Post-priming treatments that restored longevity in primed impatiens and tomato seeds were not accompanied by consistent changes in RFO or planteose contents. Our data do not rule out a role for oligosaccharides in seed longevity, but they make it unlikely that changes in oligosaccharide contents alone are responsible for the reduction in longevity due to priming or its restoration by post-priming treatments.
Nutrigenomics is the study of how constituents of the diet interact with genes, and their products, to alter phenotype and, conversely, how genes and their products metabolise these constituents into nutrients, antinutrients, and bioactive compounds. Results from molecular and genetic epidemiological studies indicate that dietary unbalance can alter gene–nutrient interactions in ways that increase the risk of developing chronic disease. The interplay of human genetic variation and environmental factors will make identifying causative genes and nutrients a formidable, but not intractable, challenge. We provide specific recommendations for how to best meet this challenge and discuss the need for new methodologies and the use of comprehensive analyses of nutrient–genotype interactions involving large and diverse populations. The objective of the present paper is to stimulate discourse and collaboration among nutrigenomic researchers and stakeholders, a process that will lead to an increase in global health and wellness by reducing health disparities in developed and developing countries.
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