Possible involvement of aquaporins in water uptake by pea seeds differing in quality

Using the method of room temperature phosphorescence (RTP), we divided air-dry pea (Pisum sativum L.) seeds subjected to accelerated ageing (40°C, 85% relative humidity) into three fractions: (I) high-quality seeds, (II) weakened seeds, and (III) dead seeds. In the process of ageing, seed germinability firstly decreased and then increased due to so-called “improved” seeds of fraction II, which returned to fraction I as judged from the RTP level; the germinability of these seeds became equal to that of fraction I seeds. Seeds capable of germination (fractions I and II) differed in the rates of imbibition, which depended on plasma membrane permeability (opened or closed water channels) but not on the presence of the seed coat. A low activation energy of seed imbibition in fraction II (less than 5 kcal/mol) indicates that water channels are open. A mercury-containing compound (5 μM p-chloromercuribenzoate (PCMB) reduced the rate of water uptake by these seeds, and dithiothreitol restored it. A high activation energy of fraction I seed imbibition (more than 12 kcal/mol) corresponded to the water uptake mainly across the lipid bilayer when water channels are closed. PCMB did not affect the rate of fraction I seed imbibition. We supposed that mature air-dry pea seeds had open water channels. During the first stages of fraction I seed imbibition, these channels were closed, limiting water uptake. NaF (100 μM), an inhibitor of phosphatase, prevented channel closing and accelerated the imbibition of fraction I seeds. It did not affect the imbibition rate of fraction II seeds, indicating their water channels to be opened. However, NaF did not affect the water uptake of “improved” fraction II seeds as well. It seems likely that their channels were closed during accelerated ageing but otherwise than via dephosphorylation. The results obtained indicate the possibility of water inflow regulation in the weakened seeds via the state of aquaporins, which form water channels in the membranes.     

The disruption of aquaporin function in cell membranes as a cause of changes in germinability of pea seeds after exposure to low doses of gamma-radiation

Pea seeds (Pisum sativum L.) from the seed lot with 80% germinability were separated in fractions according to room temperature phosphorescence: strong seeds were assigned to fraction I, and weak seeds formed to fraction II. During imbibition, the seeds of fraction II exhibited twofold higher rates of water uptake and experienced hypoxia. Some of these seeds suffocated from hypoxia, and other seeds produced seedlings with morphological defects (such seeds were considered incapable of germination). One week after irradiation with the dose of 3 Gy, germination percentage decreased to 45%, which was caused by the increase of number of weak seeds. The germinability of seeds subjected to gamma-irradiation at doses of 7 and 10 Gy was similar to that of control seeds. In these sub-lots, there appeared so-called "improved" seeds, which were similar to non-irradiated seeds in terms of phosphorescence level, the rate of water uptake and germination percentage. It was shown with the use of PCMB that the difference in the rates of water uptake by seeds of fraction I and II depended on the permeability of cell membranes. The permeability was determined by the state of aquaporins ("open"-"closed"). The experiments with phosphatase inhibitor (NaF) shown that in seeds irradiated with dose of 3 Gy (fraction II), the mechanism of aquaporins closing was broken (phosphatase was inactivated). In "improved" seeds (after irradiation with dose of 10 Gy), aquaporins were closed irreversibly in air-dry state, when aquaporin dephosphorylation was unlikely. It was concluded that the abnormal increase (following the initially decrease) in germination of pea seeds after irradiation can be explained without invoking the hypothesis on hyper function of reparatory mechanism of at low doses of irradiation.     

Expression and inhibition of aquaporins in germinating Arabidopsis seeds

Extensive and kinetically well-defined water exchanges occur during germination of seeds. A putative role for aquaporins in this process was investigated in Arabidopsis. Macro-arrays carrying aquaporin gene-specific tags and antibodies raised against aquaporin subclasses revealed two distinct aquaporin expression programs between dry seeds and young seedlings. High expression levels of a restricted number of tonoplast intrinsic protein (TIP) isoforms (TIP3;1 and/or TIP3;2, and TIP5;1) together with a low expression of all 13 plasma membrane aquaporin (PIP) isoforms was observed in dry and germinating materials. In contrast, prevalent expression of aquaporins of the TIP1, TIP2 and PIP subgroups was induced during seedling establishment. Mercury (5 microM HgCl(2)), a general blocker of aquaporins in various organisms, reduced the speed of seed germination and induced a true delay in maternal seed coat (testa) rupture and radicle emergence, by 8-9 and 25-30 h, respectively. Most importantly, mercury did not alter seed lot homogeneity nor the seed germination developmental sequence, and its effects were largely reversed by addition of 2 mM dithiothreitol, suggesting that these effects were primarily due to oxidation of cell components, possibly aquaporins, without irreversible alteration of cell integrity. Measurements of water uptake in control and mercury-treated seeds suggested that aquaporin functions are not involved in early seed imbibition (phase I) but would rather be associated with a delayed initiation of phase III, i.e. water uptake accompanying expansion and growth of the embryo. A possible role for aquaporins in germinating seeds and more generally in plant tissue growth is discussed.     

Reversal of the effects of aging in soybean seeds

Accelerated aging predisposed seeds to imbibition injury. Slowing the rate of hydration prevented the loss of germinability due to imbibition injury. Germinability of accelerated aged seeds (50 hours) was increased from 10 to 90% by controlling the rate of imbibition. Slow hydration also prevented seed electrolyte leakage. This may indicate that cell membrane permeability or rupture was a major factor contributing to the loss of germinability after aging.

Reversal of the effects of aging (repair) was accomplished by slowly inbibing and then redrying seeds (priming). This treatment lowered steep water conductivity by a factor of 2 to 5. Priming also increased the per cent germination of low vigor seeds. The mechanism of this reversal was probably metabolic because it depended on temperature, seed moisture, and treatment duration.

Priming doubled the survival of seeds in the accelerated aging vigor test. The `rejuvenation' was accepted as evidence for metabolic repair. Since the `vigor' of seeds was increased by priming, metabolic repair probably included other subcellular components as well as the plasma membrane.

     

SEEDS OF KOSTELETZKYA VIRGINICA (MALVACEAE): THEIR STRUCTURE,GERMINATION, AND SALT TOLERANCE. I. SEED STRUCTURE AND GERMINATION

Dormancy of Kosteletzkya virginica (L.) Presl. seeds is primarily due to the impermeability of the seed coat to water. The impermeable structure is assumed to be, in other Malvaceae, the palisade layer of the seed coat. The percentage of seeds capable of imbibition and germination increased with increasing time of storage at low temperatures, but the release from dormancy was not accompanied by decreased seed coat resistance to pressure. Under natural conditions, mechanical damage to the seed coat due to changes in temperature and/or abrasion may render the seeds water permeable. It is not clear what causes water permeability during storage under laboratory conditions. During seed maturation and drying, the inner epidermis of the tegmen partly separates from the rest of the seed coat and an air space, which makes the seed buoyant, is formed around the region of the chalazal cleft. The optimal temperature for germination of K. virginica seeds is between 28 and 30 C in light or darkness.     

The Importance of Genetically Determined Seed Coat Characteristics to Seed Quality in Grain Legumes

Comparisons of five pairs of isogenk lines of peas, differingonly in the A gene for seed coat colour showed that white seeds(genotype aa) imbibed more rapidly than coloured seeds (AA),suffered greater imbibition damage revealed by dead tissue onthe cotyledons, and higher solute leakage. Seed-coat pigmentationwas closely associated with slow water uptake, since when expressionof the A gene was suppressed by the recessive pollens gene,the resulting white seeds {palpal AA) imbibed rapidly. The slowwater uptake by coloured seeds was not due to the restrictionof water entry by the seed coat since the differences in imbibitionrate were maintained when a portion of the seed coat was removedand seeds were imbibed with the exposed cotyledon in contactwith moist filter paper. Imbibition of similarly treated seedsby immersion in polyethylene glycol solutions (1–4%) whichincreased the seed/solution wettability, had little effect onthe water uptake of coloured seeds compared to imbibition inwater whereas that of white seeds increased in the first 10mins imbibition. Poor wettability of the inner surface of colouredseed coats did not therefore explain the slow imbibition ofthese seeds. The white seed coats loosened rapidly during imbibitionwhilst the coloured seed coats remained closely associated withthe cotyledons suggesting that the adherence of the seed coatto the cotyledons and therefore the ease of access of waterbetween the testa and cotyledons determines the rate of imbibition.The rapid water uptake by white-coated seeds and the subsequentimbibition damage may explain the high incidence of infectionof these seeds by the soil-bome fungus Pythhan after 2 d insoil. Improved seed quality and emergence may therefore be achievedby breeding for seed coat characteristics leading to reducedrates of imbibition Pisum sativum, isogenic lines, A gene, seed coat colour, imbibition, imbibition damage, wettability, pollens gene, seed quality, grain legumes     

Allelopathy due to purine alkaloids in tea seeds during germination

Summary During imbibition of whole tea seeds (6 days) two purine alkaloids, caffeine and theobromine, did not decrease in the seed coats and there was no increase in the seeds. In parallel with and after the breaking of seed coats there was a gradual release of caffeine from coats of germinating seeds. By contrast, when the seed was freed from the outer seed coat and soaked, imbibition of the seed required only 2 days and simultaneously caffeine was released from the inner seed coat. In such seeds, but not in whole seeds, growth of embryonic tissues (roots and shoots) was inhibited after the breaking of the inner seed coats. Nevertheless, caffeine increased more in such roots of the seedlings of decoated seeds than in roots of normal seedlings.     

Seed coat structure and oxygen availability control lowtemperature germination of melon (Cucumis melo) seeds

The involvement of the seed coat in low-temperature germination of melon seeds was examined in two accessions differing in their ability to germinate at 14°C: Noy Yizre'el (a cold-sensitive cultivar) and Persia 202 (a cold-tolerant breeding line). Decoating resulted in full germination of Noy Yizre'el at 14°C, but splitting the coat increased germination only partially. Thus, the inhibition of Noy Yizre'el germination at 14°C is not due to physical constraint on radicle protrusion. At 25°C, seeds of both accessions submerged in water or agar germinated fully as long as the hilum aperture remained uncovered. Submerging the whole seed, or covering the hilum with lanolin, strongly depressed germination of Noy Yizre'el but not of Persia 202. Accessions differed in germination response to decreasing O₂ concentration, with Noy Yizre'el showing higher sensitivity to hypoxia. These differences were correlated with differences in seed coat structure as well as in embryo sensitivity to hypoxia. Intercellular spaces in the outer layer of the seed coat were evident in the more tolerant Persia 202, while in the sensitive Noy Yizre'el this layer was completely sealed. Sensitivity to hypoxia increased at 15°C as compared with 25°C, the increase being greater in Noy Yizre'el. It is proposed that the seed coat-imposed dormancy at low temperature in Noy Yizre'el is the combined result of more restricted oxygen diffusion through the seed coat and a greater embryo sensitivity to hypoxia, rather than to physical constraints of radicle break-through or impairment of imbibition.     

Bimodal changes in germinability of pea seeds under the influence of low doses of gamma-radiation

Pea seeds (cv. 'Nemchinovskii-85', harvest of 2002, 80%-germination percentage) were exposed to gamma-radiation with doses ranging from 19 cGy to 100 Gy. One week after the irradiation with doses of 19 cGy and 3 Gy. the germination percentage decreased to 58 and 45%, respectively; at doses of 7 and 10 Gy it was 73 and 70% respectively. At greater doses (25, 50, and 100 Gy), germination percentage decreased in proportion. Anomalous changes in seed germination percentage (as a function of irradiation dose) were caused by the redistribution of irradiated seeds between fractions I and II. The measurements of room temperature phosphorescence in air-dry seeds and the phosphorescence of endogenous porphyrines of imbibing seeds showen that the germination decrease after the irradiation with low doses (19 cGy and 3 Gy) was caused by the increase in the number of weak seeds of fraction II, which had high rates of water uptake and suffered from hypoxia under seed coat. Some of these seeds suffocated from hypoxia, and other seeds produced seedlings with morphological defects (such seeds were considered incapable of germination). During storage of seeds irradiated at doses 19 cGy-10 Gy, the recovery of germination percentage (after its initial decrease) was caused by the decrease in seed number in fraction II. The subsequent germination decrease was caused by seed death. The higher was the irradiation dose, the faster were changes in germination percentage during storage of irradiated seeds. Bimodal changes in pea seed germination with the increase of y-irradiation dose has apparently the same origin as the changes in seed germination during accelerated ageing.     

The effect soaking pea seeds with or without seedcoats has on seedling growth

Pea seeds (Pisum sativum L. `Alaska') with intact seedcoats (WC) and with seedcoats removed (WOC) were soaked in distilled water for 24 hours at 20°. The water, containing the pea diffusate, was decanted after the second, fourth, sixth, eighth, twelfth, and twenty-fourth hour and analyzed for total nitrogen, α-amino nitrogen, carbohydrate, and total solute dry weight. The seeds were germinated at 20° in a 16 hour photoperiod of 300 foot candles. Stem lengths and dry weights of roots, shoots and cotyledons were determined after 4, 11, and 18 days of growth. WOC seeds imbibed more water than WC seeds during the 24 hour imbibition period. Diffusates from WOC seeds always contained more solute than diffusates from WC seeds. Maltose, glucose, and fructose were not detected in the early diffusates from WOC seeds but were found in WC seed diffusates at all times. Seedlings from WC seeds had longer stems than those from WOC seeds. The dry weight of stems and roots of WC seedlings was greater than those from WOC seedlings. The dry weight of cotyledons from 18 day-old WC seedlings was less than from WOC seedlings. Water absorption by WC seeds was slower than by WOC seeds. Removal of the seedcoat allowed rapid imbibition resulting in seed injury presumably because of the loss of solutes which included monosaccharides, disaccharides, amino acids, and other nitrogen containing compounds. These results are consistent with the hypothesis that rapid imbibition disrupts membrane organization leading to reduction of seedling growth.     

MORPHOLOGICAL OBSERVATIONS ON THE EARLY IMBIBITION OF WATER BY SIDA SPINOSA (MALVACEAE) SEED

The chalazal area is confirmed as the site of initial water entry into prickly sida (Sida spinosa L.) seeds. Very early during imbibition of water, a kidney-shaped area of the seed coat separates from underlying cells forming a blister. This blister may also be induced in dry seeds (both afterripened and nonafterripened) when pressure is applied to the chalazal area. Blisters form more readily on afterripened seeds than on nonafterripened seeds, however, and the event is correlated with an increase in seed coat permeability to water. Immediately beneath the palisade layer of the blister lies a single layer of subpalisade cells. This layer is observed only in the region of blister formation. As the blister separates, the end walls of the subpalisade cells remain attached to the floor of the palisade layer. The subpalisade cells are thereby broken open, and their contents disgorged into the blister lumen. Evidence indicates that this separation of the palisade and subpalisade layers in the chalazal area initiates imbibition of water by prickly sida seeds.     

金钟藤种子低萌发率原因探讨

金钟藤种子在室内萌发率很低,为进一步探讨金钟藤种子的特性,阐明其种子萌发率低的主要原因,对金钟藤种皮的透水性、种子解剖结构、种子活力和种子内源抑制物的生物测定进行了研究。结果表明:金钟藤种皮透水性较差,完整种子比破皮种子吸水达到最高水平慢38h;种子空瘪粒多,占所有种子的30%;种子活力较低,平均活力仅为35%;金钟藤种子甲醇粗提液对白菜种子萌发率、根长和芽长均有较强的抑制作用,其浸提液浓度在25mg/mL时,严重抑制白菜种子萌发和生长,即金钟藤种子内部含有较高的内源抑制物质。金钟藤种子萌发率低,表明其近年来突发性蔓延成灾主要不是由种子生成新个体造成的,导致其蔓延成灾的关键因素还需要进一步深入研究。     

Lipid peroxidation, carbohydrate hydrolysis, and Amadori-Maillard reaction at early stages of dry seed aging

Processes underlying aging of air-dry seeds were investigated. Naturally aged seeds of pea (Pisum sativum L.), cucumber (Cucumis sativus L.), and buckwheat (Fagopyrum esculentum Moench.) were separated into three fractions of different quality according to their room temperature phosphorescence (RTP): fraction I contained strong seeds; fraction II comprised weak seeds; and fraction III was composed of dead seeds. These seed fractions were used to identify the processes in air-dry seeds that account for the initial deterioration in seed quality at early stages of aging and for the possible transient improvement of seed quality at further aging. Experiments with seed powders showed that the increase in thermochemiluminescence (TCL) in the temperature range of 50?C110°C was determined by the presence of lipid peroxidation products. No difference was observed between TCL levels in fraction I and fraction II seeds; hence, lipid peroxidation is not responsible for the transition of strong seeds to the fraction of weak seeds. The TCL intensity monitored upon heating in the range of 50?C110°C increased only in the fraction of dead seeds. In fraction II seeds, a twofold increase in TCL upon heating at 150°C was observed in aged seeds; this indicates that the products of oligosaccharide hydrolysis (glucose) were more abundant in fraction II seeds than in fraction I seeds under similar conditions. This assumption was confirmed by direct determination of glucose content with a glucometer. Hence, the transition of air-dry seeds from fraction I (strong seeds) to fraction II (weak seeds) occurred due to the activation of carbohydrate hydrolysis in aging seeds. Since air-dry seeds contain no free water during aging, the seed moisture content decreases during hydrolysis, presumably, because the bound water participates in the hydrolysis reaction. The decrease in ??improved?? seeds of glucose content and the increase in moisture content suggest the activation of amino-carbonyl reaction. It is proposed that the amino-carbonyl (Amadori-Maillard) reaction is responsible for closing water channels in improved seeds, as well as for the decreased water permeability of cell membranes during seed imbibition.     

Imbibition period as the critical temperature sensitive stage in germination of lima bean seeds

Lima bean seeds (Phaseolus lunatus L.) and excised embryonic axes can be injured during imbibition at temperatures below 25°. The early imbibitional stage is critical; imbibition at 25° followed by low temperature exposure does not cause injury. Sensitivity to chilling injury is conditioned by the pre-harvest seed history. Low vigor (bleached) seeds are most sensitive to injury, the effects of which can be intensified by restricted oxygen supply during early axis growth. The seed coat, by preventing water uptake, can permit the seed to avoid injury. This protective mechanism is most effective at low temperature and high moisture stress. Immediately following low temperature imbibition, injured axes lose organic materials, probably nucleotides. This organic leachate is a potential influence on soil microorganisms and, together with the temperature sensitivity, vigor, and seed coat effect undoubtedly is important in controlling the potential variability in germination shown by a seed population.     

Germination and Food Reserves in Bambarra Groundnut Seeds (Voandzeia subterranea Thouars) after Different Periods of Storage

Studies were made on bambarranut seeds (Voandzeia subterraneaThouars) after 0, 6, 12, 18 and 24 months of storage in gunnybags under laboratory conditions (25–35 °C). Seeddeterioration during storage was indicated by delayed germination,reduced germinability, reduced growth of seedlings and increasednumber of stunted seedlings culminating in a total failure ofgermination after two years. Slight depletion of food reserves occurred during seed storage.The loss in fat was higher than starch or protein. Total solublesugars decreased while the content of total fatty acids andamino acids and soluble protein increased. Total nitrogen (N)remained unaffected while soluble-N and amino-N increased. Allthese components showed a rapid change (increase or decrease)from 12 months to 18 months of storage which was associatedwith commencement of rapid decline in germinability of the seedsand growth of the seedlings. Initial rapid imbibition of water was observed in viable aswell as non-viable seeds, though at a higher rate in the latterand followed by a lag period in both. At the end of 24 h ofimbibition, water content in non-viable seeds was less thanthat in viable ones. Key words: Voandzeia subterranea, Seed germination, Seed storage     

Acquisition of physical dormancy and ontogeny of the micropyle--water-gap complex in developing seeds of Geranium carolinianum (Geraniaceae)

Background and Aims

The ‘hinged valve gap’ has been previously identified as the initial site of water entry (i.e. water gap) in physically dormant (PY) seeds of Geranium carolinianum (Geraniaceae). However, neither the ontogeny of the hinged valve gap nor acquisition of PY by seeds of Geraniaceae has been studied previously. The aims of the present study were to investigate the physiological events related to acquisition of PY and the ontogeny of the hinged valve gap and seed coat of G. carolinianum.

Methods

Seeds of G. carolinianum were studied from the ovule stage until dispersal. The developmental stages of acquisition of germinability, physiological maturity and PY were determined by seed measurement, germination and imbibition experiments using intact seeds and isolated embryos of both fresh and slow-dried seeds. Ontogeny of the seed coat and water gap was studied using light microscopy.

Key Results

Developing seeds achieved germinability, physiological maturity and PY on days 9, 14 and 20 after pollination (DAP), respectively. The critical moisture content of seeds on acquisition of PY was 11 %. Slow-drying caused the stage of acquisition of PY to shift from 20 to 13 DAP. Greater extent of cell division and differentiation at the micropyle, water gap and chalaza than at the rest of the seed coat resulted in particular anatomical features. Palisade and subpalisade cells of varying forms developed in these sites. A clear demarcation between the water gap and micropyle is not evident due to their close proximity.

Conclusions

Acquisition of PY in seeds of G. carolinianum occurs after physiological maturity and is triggered by maturation drying. The micropyle and water gap cannot be considered as two separate entities, and thus it is more appropriate to consider them together as a ‘micropyle–water-gap complex’.     

Aqueous pathways dominate permeation of solutes across Pisum sativum seed coats and mediate solute transport via diffusion and bulk flow of water

The permeability of seed coats to solutes either of biological or anthropogenic origin plays a major role in germination, seedling growth and seed treatment by pesticides. An experimental set‐up was designed for investigating the mechanisms of seed coat permeation, which allows steady‐state experiments with isolated seed coats of Pisum sativum. Permeances were measured for a set of organic model compounds with different physicochemical properties and sizes. The results show that narrow aqueous pathways dominate the diffusion of solutes across pea seed coats, as indicated by a correlation of permeances with the molecular sizes of the compounds instead of their lipophilicity. Further indicators for an aqueous pathway are small size selectivity and a small effect of temperature on permeation. The application of an osmotic water potential gradient across isolated seed coats leads to an increase in solute transfer, indicating that the aqueous pathways form a water‐filled continuum across the seed coat allowing the bulk flow of water. Thus, the uptake of organic solutes across pea testae has two components: (1) by diffusion and (2) by bulk water inflow, which, however, is relevant only during imbibition.     

Seed Coat Dormancy in Two Species of Grevillea(Proteaceae)

The role played by the seed coat in seed dormancy of Grevillealinearifolia(Cav.) Druce and G. wilsonii(A. Cunn.) was testedby a series of manipulations in which the seed coat was dissectedand removed, dissected and returned to the decoated seed, ordissected, removed and given a heat shock, and returned to thedecoated seed. Germination of intact seeds of both species wasalso examined after exposure to heat shock, smoke, or heat shockand smoke combined. Water permeability of the seed coat wasinvestigated by examining imbibition. For intact seeds, virtuallyno germination occurred under any treatment (G. wilsonii), orgermination was increased by exposure to either heat or smoke(G. linearifolia). Removal of the seed coat led to germinationof all decoated seeds for G. linearifolia, or a proportion ofdecoated seeds for G. wilsonii. Inclusion of smoked water inthe incubation medium led to a higher proportion of decoatedseeds germinating for G. wilsonii. Returning the seed coat,either with or without heat shock to the seed coat, did notsignificantly affect germination in either species. Seed coatswere permeable to water in both species. For the two Grevilleaspecies, there were different dormancy mechanisms that werecontrolled by the seed coat (G. linearifolia) or by both theseed coat and embryo (G. wilsonii). Copyright 2000 Annals ofBotany Company Grevillea linearifolia, Grevillea wilsonii, dormancy, seed coat dormancy, seed coat permeability, smoke, heat shock, germination     

The role of the persistent fruit wall in seed water regulation in Raphanus raphanistrum (Brassicaceae)

Background and Aims

Dry fruits remain around the seeds at dispersal in a number of species, especially the Brassicaceae. Explanations for this vary, but usually involve mechanisms of innate dormancy. We speculate that, instead, a persistent fruit may give additional protection through control of dehydration, to species growing in arid or Mediterranean environments where water is sporadic.

Methods

X-rays and weight measurements were used to determine the extent to which Raphanus raphanistrum seeds within mature fruits imbibe water, and germination tests determined the roles of the fruit and seed coat in seed dormancy. Rates of water uptake and desiccation, and seedling emergence were compared with and without the fruit. Finally, germinability of seeds extracted from fruits was determined after various periods of moist conditions followed by a range of dry conditions.

Key Results

Most seeds rapidly take up water within the fruit, but they do not fully imbibe when compared with naked seeds. The seed coat is more important than the dry fruit wall in maintaining seed dormancy. The presence of a dry fruit slows emergence from the soil by up to 6–8 weeks. The fruit slows the rate of desiccation of the seed to a limited extent. The presence of the fruit for a few days during imbibition somehow primes more seeds to germinate than if the fruit is absent; longer moist periods within the pod appear to induce dormancy.

Conclusions

The fruit certainly modifies the seed environment as external conditions change between wet and dry, but not to a great extent. The major role seems to be: (a) the physical restriction of imbibition and germination; and (b) the release and then re-imposition of dormancy within the seed. The ecological significance of the results requires more research under field conditions.     

Functional aspects of mature seed coat of theCannaceae

Cannaceae seeds have been analysed regarding seed coat structure, germination and macromolecular composition of the seed coats. Data of several mass spectrometric techniques were combined with those of microscopic and histochemical techniques to acquire insight into the functions of the seed coat.Cannaceae seeds have an exotestal layer of Malpighian cells with a hydrophobic and a hydrophilic part. The hydrophobic part is mainly responsible for the impermeability of the seed and contains silica, callose, lignin as water repellent substances. Water can only enter the seed after a certain temperature-induced opening of an imbibition lid. During imbibition the hydrophilic part of the Malpighian cells swells and the seed coat ruptures due to differences in pressure in the upper and lower part of the Malpighian cells.