Development of pepper (Capsicum annuum) seed quality

Changes in seed quality in pepper (Capsicum annuum L.) were monitored during seed development and maturation in two seasons. Seed quality was assessed by a number of different tests, but principally by determining seed storage longevity in laboratory tests and seedling growth in glasshouse tests. Mass maturity (defined as the end of the seed-filling phase) occurred 49–53 days after anthesis (DAA) in 1989 (varying among fruit layers) and 53 DAA in 1990 when seed moisture contents were 51–53%. The onset of both germinability and desiccation tolerance occurred either just before or at mass maturity. Maximum potential longevity (assessed by the value of the seed lot constant K_i) was achieved 63–65 DAA, i.e. not until 10–12 days after mass maturity (DAMM), in both years. Seedling dry weights in the glasshouse growth tests were maximal later still - for seeds harvested 17–21 DAMM in 1989 and 17 DAMM in 1990; the effects on seedling weight arose from differences in times from sowing to emergence (P < 0.005) among different seed harvests, with no significant differences in subsequent relative growth rates (P > 0.25). Seed priming reduced mean germination times for seeds harvested at all stages of development, but had little effect on germination capacity and potential longevity, and did not affect the pattern of changes in potential longevity during seed development and maturation. The results contradict the hypothesis that seed quality is maximal at the end of the seed-filling phase and that viability and vigour begin to decline immediately thereafter.     

Relationships between indices of seed maturity and carrot seed quality

Carrot seeds cvs Chantenay and Amsterdam were harvested on several occasions from crops grown in 1985 and 1986, from the time they had reached or were close to their maximum dry weight and were starting to turn brown. Maximum seed dry weight occurred approximately 40–45 days after flowering (DAF) in both cultivars. Maximum germination (International Seed Testing Association 14-day count) occurred 40 and 55 DAF in cvs Chantenay and Amsterdam, respectively, but the maximum 7-day count and the minimum coefficient of variation of embryo length did not occur until 60 DAF in cv. Chantenay and 55 to 65 DAF in cv. Amsterdam. Percentage germination was negatively and linearly related to seed moisture content, chlorophyll a + b content in the seed coat and seed distortion, measured on a modified tensile-testing machine, the relationships accounting for 77, 71 and 64% of the variance in the 7-day germination count, respectively. The corresponding values for the 14-day count were 63, 61 and 50%. A simple field test for monitoring seed maturity is proposed.     

Self- and open-pollination as factors influencing seed quality in leek (Allium porrum)

The effect of self-pollination compared with open-pollination on leek seed weight, and subsequent seedling emergence and growth were observed for two years of seed production. Open-pollination gave higher mean seed weights and lower seedling losses during plant establishment than self-pollination. Depending upon the year, the individual plant and pollination method, between 2. 4 and 56.6% of the variation in seedling weight c. 4–5 wk after sowing under controlled conditions was accounted for by differences in seed weight and time of seedling emergence. In 1984 but not 1983, a greater proportion of the variation in seedling weight was accounted for by seed weight and time of seedling emergence in open than self-pollinated seeds.     

Interactions between seed priming treatments and nine seed lots of carrot, celery and onion. II. Seedling emergence and plant growth

Samples of three seed lots of each of three cultivars of carrot, celery and onion were primed in polyethylene glycol solution for two weeks at 15 °C. Seedling emergence was recorded in the field for carrot and onion and in the glasshouse for celery. Compared to the untreated control, priming increased the percentage seedling emergence in certain poorly-emerging seed lots of carrot and celery, but had no effect on onion. Mean emergence times were reduced by priming in all seed lots, by 3–5, 5–8 and 3–9 days in carrot, celery and onion respectively. The largest effects occurred in the slowest-emerging seed lots. There were significant interactions between priming and seed lots within cultivars in carrot and celery and between priming and cultivars in celery and onion. Priming generally reduced the spread of emergence times, but the effects were not statistically significant in carrot. Drying back the primed seeds had no effect on percentage emergence in onion, but reduced it (compared to primed seed which had not been dried-back) in certain carrot and celery seed lots. Primed and dried-back seeds emerged later than primed seeds, by up to 1·5, 2·6 and 2·6 days in carrot, celery and onion respectively. The spread of emergence times was generally larger for primed and dried-back seeds than for primed seeds, but the differences were not always statistically significant. Plant fresh weights were recorded 9, 7 and 12 wk after sowing for carrot, celery and onion, respectively. In each species, mean plant weight was inversely related to seedling emergence time; thus plants grown from primed seed were always heavier than the controls, by up to 33%, 182% and 47% in carrot, celery and onion respectively.     

A comparison of methods for evaluating seed quality in carrots (Daucus carota)

Comparisons of embryo length, ISTA (International Seed Testing Association), cold (10 ^oC), controlled-deterioration, and slope tests for estimating seedling size variability and percentage seedling emergence were made using eighteen stocks of carrot seeds. The coefficient of variation (C.V.) of seedling weight was closely correlated with the C.V. of embryo length (r = 0–87 D.F., 16) and also with the C.V. of root length from the slope test (r = 0–63 D.F., 16). Poorer estimates of percentage seedling emergence were obtained from the cold test in all three sowings (correlation coefficients were between 0–65 and 0–70 D.F., 16) compared with the ISTA, controlled deterioration, and slope tests. For these three tests the correlation coefficients varied between 0–80 and 0–87 (D.F., 16) for all sowings. The embryo test is a rapid (1 man hour per seed lot) and accurate method of assessing potential variability between seed lots. The slope test could provide a less labour demanding method of assessing potential variability than the embryo length test though the results would not be available immediately. However, the slope test could also provide as good an estimate of seedling emergence in the field as the ISTA or controlled-deterioration tests.     

Priming of field-sown rice seed enhances germination, seedling establishment, allometry and yield

Poor seedling establishment is a major deterrent in adopting direct seeding of rice. Seed priming to obtain better crop stand could be an attractive approach. The objective of this study was to determine the effectiveness of seed priming strategies on the improved agronomic characters of direct-sown rice. Seed priming strategies were: hydropriming for 48 h, osmohardening with KCl or CaCl₂ for 24 h, ascorbate priming for 48 h and seed hardening for 24 h, pre-germination (traditional soaking for nursery raising) and untreated control. Seed priming improved germination and emergence, allometry, kernel yield, and its quality, whilst pre-germination displayed poor and erratic emergence of seedling followed by poor plant performance. Faster and uniform emergence was due to improved α-amylase activity, which increased the level of soluble sugars in the primed kernels. Osmohardening with KCl gave greater kernel and straw yield and harvest index, followed by that of CaCl₂, hardening and ascorbate priming. Improved yield was attributed principally to number of fertile tillers and 1000 kernel weight. A positive correlation between mean emergence time and days to heading, while a negative one between kernel yield and harvest index suggested long-term effects of seed priming on plant growth and development. The results suggest that physiological changes produced by osmohardening enhanced the starch hydrolysis and made more sugars available for embryo growth, vigorous seedling production and, later on, improved allometric, kernel yield and quality attributes.     

Non-lethal freezing effects on seed degreening in Brassica napus

The effects of a non-lethal freezing stress on chlorophyll content, moisture level and distribution, and abscisic acid (ABA) levels were examined in siliques and seeds of Brassica napus (canola). A non-lethal freezing stress resulted in the retention of chlorophyll in seed at harvest that was most pronounced for seeds 28, 32 and 36 days after flowering (DAF). This increase was primarily due to an increased retention of chlorophyll a relative to chlorophyll b. Chlorophyll retention in seeds exposed to a non-lethal freezing stress correlated with an increased ABA catabolism, as measured 1, 3 or 7 days after the stress treatment. Although the non-lethal freezing stress had no significant effect on moisture content in seeds of siliques stressed at 28–44 DAF, moisture distribution, as viewed by magnetic resonance imaging, showed an uneven drying of 32 and 40 DAF siliques after exposure to the non-lethal freezing stress. Moisture was initially lost more rapidly from the silique wall between seeds, than in control non-stressed siliques. Increased moisture loss was not due to structural changes in the vasculature of the silique/seed of stressed tissues. These results are consistent with the hypothesis that a non-lethal freezing stress-induced decrease in ABA level, during seed maturation, effects an inhibition of normal chlorophyll a catabolism resulting in mature but green B. napus seed.     

Effects of Planting Depth,Sediment Grain Size,and Nutrients on Ruppia maritima and Potamogeton perfoliatus Seedling Emergence and Growth

Protocols are now available for seed harvest, storage and germination of several mesohaline and polyhaline species; however, low seedling survival rates point to the need for an increased understanding of factors affecting seedling establishment. Depth of seed burial in sediments and initial seedling growth rates are shown to be limiting factors for photosynthetic competency of Ruppia maritima and Potamogeton perfoliatus. Seedling emergence is inversely proportional to planting depth on sediments ranging in grain size from coarse sands (850 μm) to silt (63 μm). Less than 6% of the seeds of either species emerged when buried to a depth of 3 cm in test sediments. Germination was greatest for seeds placed on the surface of sediments; however, these seedlings were subject to displacement because of the weak and fragile roots produced during early growth. Fine sediments may be more favorable for R. maritima seedling establishment, because seedling emergence and height decreased with increasing sediment grain size. Potamogeton perfoliatus seedlings seem to be more tolerant of a wider range of sediment grain sizes than R. maritima as indicated by the lack of an effect of sediment grain size on P. perfoliatus seed emergence, seedling height, and biomass. Increasing nutrients stimulated seedlings of both species; however, even at the highest concentrations tested, growth, as determined by shoot elongation and leaf and root formation, slowed within 7–10 days. This suggests factors other than mineral nutrients and light limit growth or that growth shifts from aboveground biomass production to belowground vegetative spread.     

The effect of plant density, harvest date and method on the yield of seed and components of yield of parsnip (Pastinaca sativa)

Experiments conducted over four years (1980–1983) with parsnip cv. Avonre-sister examined the effects of plant density, harvest date and method on seed yield and the components of yield. In 1980, using the root-to-seed method, the yield of seed increased by 50% for an increase in plant density from 5.6 to 35 plants m^-2; the maximum yield at the optimum harvest date was 3015 kg h^-1. In the other three years, using the seed-to-seed method, seed yield increased to a maximum with an increase in plant density from c. 1 to 10 plants m^-2 but at higher densities yields were lower. At c. 10 plants m^-2, seed yields were 1567, 4029 and 1040 kg ha^-1, in 1981, 1982 and 1983, respectively, when harvested at the optimum time. An increase in plant density increased the number of non-seeding plants in the population, reduced the number of umbels per plant and reduced the number of seeds per umbel. Mean seed weight was, in general, unaffected by plant density and so effects of density on yield reflected changes in seed numbers. Increasing the plant density increased the proportion of primary-umbel seed. Maximum yields of seed were obtained from crops harvested at a seed-moisture content of 50 to 70%, which occurred 46 ± 2·4 days after flowering. Delay in harvesting after this date led to a loss of yield of 33, 35, 139 and 32 kg ha^-1wk^-1 in 1980, 1981, 1982 and 1983, respectively. Plants cut, placed in the windrow to dry and then threshed gave similar yields to those harvested at the same time but dried in bins.     

A Comparison of Leek (Allium porrum) and Onion (Allium cepa) Seed Development

Leek and onion seed dry weight increased exponentially for thefirst 31 days after flowering (DAF) but thereafter the increasein dry weight was slower. Before maximum seed dry weight wasreached at 45 DAF in onion and 59 to 66 DAF in leek, seed moisturecontent, seed oxygen uptake and conductivity of the seed steepwater fell from initially high levels. Although some seeds germinated31 DAF in both species, full germination in both was not reacheduntil 66–80 DAF. Tolerance of the seed to artificial dryingimmediately after harvesting occurred 45 DAF in onion and 74DAF in leek. Free nuclear division continued in the endospermuntil 17–22 DAF in onion and until 31–35 DAF inleek but it was not until 45 DAF in onion and 66 DAF in leekthat the embryo and endosperm filled the cavity formed by thepericarp. After formation of cell walls in the endosperm thepattern of change in cell number in both species was similar.The shrunken appearance of the seed coat in leek, which occurredearly in seed development, was associated with the period offree nuclear division in the endosperm and, in addition, thepericarp was thinner than in onion. There was no evidence thatthe shrunken seed coat early in development was associated withself as opposed to open-pollination. Allium porrum, Allium cepa, seed development, endosperm, embryo, cell number, germination, respiration, seed leachates     

Efficiency of endozoochorous seed dispersal in six dry-fruited species (Cistaceae): from seed ingestion to early seedling establishment

We combined laboratory and nursery experiments to analyse the effectiveness of sheep as endozoochorous seed dispersers of six native shrubby Cistaceae species collected in SE Spain (Helianthemum apenninum (L.) Mill., H. violaceum (Cav.) Pers., Fumana ericoides (Cav.) Grand., F. thymifolia (L.) Spach, Cistus monspeliensis L. and C. laurifolius L.), considering the main stages after seed ingestion, i.e. seed recovery, seed germination, seedling emergence and early seedling establishment. Seed recovery after gut passage was high (around 40%) for all the species, except F. thymifolia (12%). Most seeds (ca. 90%) were recovered within 48 h after ingestion for all the species, although seeds were still recovered up to 96 h after ingestion. Gut passage increased germination up to seven-fold compared to non-ingested seeds. Furthermore, seedling emergence from seeds contained in pellets was overall similar (intact pellets) to or higher (crumbled pellets) than emergence from seeds without dung. Survival of emerged seedlings and mass of seedlings after 20 days were not reduced by dung. Sheep act therefore as effective dispersers of these Cistaceae species by scattering seeds and promoting germination, while faeces do not hamper seedling establishment. We conclude that the interaction between herbivorous ungulates and these dry-fruited species may be considered a mutualism qualitatively similar to the mutualism between frugivorous vertebrates and fleshy-fruited plants.     

Viability of onion (Allium cega) seed as influenced by temperature during seed growth

The yield and performance of seeds from crops of winter-hardy, bolting-resistant onion grown at temperatures of 15–16, 18–19 and 22–23°C in 1979, 1980 and 1982 were compared. Yields of seed from crops grown at 22–23°C were lower than those from crops grown at lower temperatures but the seeds ripened between 11 and 32 days earlier. Seeds from crops grown at mean temperatures of above 18°C gave higher percentage germination when imbibed at 30°C than 20°C and they also gave higher percentage seedling emergence than those from crops grown at lower temperatures. Seedlings from seeds produced at mean temperatures above 18°C were heavier than those from seeds of a similar weight but produced at lower temperatures. None of these differences were associated with differences in seed weight, embryo weight or seed dormancy but were positively correlated with differences in seed N-concentration. The differences were also associated with the rate of imbibition of water as high germination, high N-content seeds had a slower rate of imbibition than low germination, low N-content seeds of the same weight.     

Influence of post-flowering temperature on seed development, and subsequent performance of crisp lettuce

Crisp lettuce plants cv. Saladin were grown from the time they started flowering, at 20/10°C (16 h day, 8 h night), 25/15°C and 30/20°C in glasshouses on two occasions in 1985. Yields of seed increased from, on average, 15 g to 27 g and then fell to 20 g per plant with progressive increases in temperature. The number of mature florets per plant increased with temperature but the number of seeds per mature floret was lower at 20/10°C and 30/20°C than at 25/15°C. An increase in temperature reduced mean seed weight by up to 45%, seed volume by 15%, cell numerical volume density (N_v) by 27% and the number of cells per seed by 39%. Percentage seed germination reached a maximum early in seed development at the stage when the pappus appeared through the involucral bracts. Differences in percentage germination and vigour of seeds (slope test) from different temperatures were accounted for largely by the effects on mean seed weight. However, when germinated at 30°C seeds produced at 30/20°C germinated more readily than those produced at 25/15°C or 20/10°C. Seed vigour gradually increased with an increase in the length of storage after harvest, reaching a maximum after 260 days. In general, seeds produced at 25/15°C exhibited a greater variation in numbers of seeds per floret, N_v, seed weight, times of seedling emergence, seedling and mature head weight than seeds produced at lower or higher temperatures.     

Increasing phosphorus concentration in seed of annual pasture legume species increases herbage and seed yields

In glasshouse experiments with low levels of soil applied phosphorus (P), yields of four annual pasture legumes (Medicago polymorpha, Trifolium subterraneum, T. balansae, Ornithopus compressus) increased with increasing P concentration in the seed. In a further experiment, M. polymorpha cv. Serena was grown at the same plant density from seed of two P concentrations and two seed sizes when two levels of finely ground superphosphate were applied to the soil. Higher P concentrations in the seed increased yields of dried tops by about 30% for the first harvest (21 days), 20% for the second harvest (52 days), and 9% at maturity (103 days), and seed yields by 11%. Larger seeds increased yields of dried tops by between 6–46% for the first two harvests but at maturity yields of dried tops and seed were unaffected by seed size. None of the interactions were statistically significant (P>0.05), except for the first harvest when two interactions (P concentration in the seed × seed size (i.e. P content in seed), and P applied to the soil × P concentration in the seed × seed size) were significant at P<0.05 level. In a field experiment, Trifolium subterraneum clover seed (two cultivars) of the same size but with two different P concentrations was sown at the same plant density and two levels of granulated (0.2–5 mm) superphosphate were applied to the soil surface. The higher level of superphosphate increased dried herbage yields of the dense clover swards by three- to four-fold 90 and 120 days after sowing. The higher P concentration in the seed increased yields of dried herbage by between 50 to 25%, depending on the level of P applied to the soil and the harvest date.     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

Effects of seed dispersal on Juniperus communis recruitment on a Mediterranean mountain

Abstract. The recruitment of the relict shrub Juniperus communis on a mountain in SE Spain was studied during the period 1994–1998. The main objective was to determine both the quantitative and qualitative effects of bird dispersal on seedling establishment. Seed removal by birds, seed rain, post‐dispersal seed predation, germination, and seedling emergence and survival were analysed in different microhabitats. Birds removed 53 ‐ 89% of the seeds produced by plants. Seed rain was spatially irregular as most seeds accumulated near stones used by birds as perches and below mother plants while a few seeds were dropped in wet meadows and open ground areas. Post‐dispersal seed predation by rodents affected < 10% of dispersed seeds but varied significantly among microhabitats. Only 3.6 ‐ 5.5% of dispersed seeds appeared viable, as many seeds had aborted or showed wasp damage. Seeds germinated in the second and third springs after sowing, reaching a germination percentage of 36%. Seedling emergence was concentrated in wet meadows. Seedling mortality was high (75–80%), but significantly lower in wet meadows, the only microhabitat where seedlings could escape from summer drought, the main mortality cause. Seed abortion, germination and seedling mortality proved to be the main regeneration constraints of J. communis on Mediterranean mountains. Birds exerted a strong demographic effect, although their qualitative effect was limited by abiotic factors which caused the pattern of seed rain to differ from the final pattern of recruitment between microhabitats.     

From seed to seedling: A critical transitional stage for the Mediterranean psammophilous species Dianthus morisianus (Caryophyllaceae)

Abstract

Seed germination, seedling emergence and seed persistence in the soil were investigated for Dianthus morisianus (Caryophyllaceae), a psammophilous endemic species of Sardinia. Stored and freshly collected seeds were incubated in a range of constant temperatures (5–25°C) and an alternating temperature regime (25/10°C). The effect of seed burial depth on seedling emergence was investigated under controlled environmental conditions. Seed persistence in the soil was verified by in situ experimental seed burials. Seeds of this species were non-dormant, and all seed lots germinated both in the light and darkness, mainly at low temperatures (≤20°C), with a maximum at 15°C (≥95%). Optimal seedling emergence was obtained when seeds were buried at a depth of 1–2 cm, and a declining emergence with increasing depth was observed. D. morisianus was also unable to form a persistent soil seed bank. The fate of the seeds that, after dispersal, do not emerge from the soil in the spring is, therefore, presumably to die before the next favourable growing season.     

Arabidopsis sucrose synthase 2 and 3 modulate metabolic homeostasis and direct carbon towards starch synthesis in developing seeds

Two genes encoding sucrose synthase (SUS), namely SUS2 (At5g49190) and SUS3 (At4g02280), are strongly and differentially expressed in Arabidopsis seed. Detailed biochemical analysis was carried out in developing seeds 9–21 days after flowering (DAF) of wild type and two knockouts. SUS2 and SUS3 are not redundant genes since single knockouts show a phenotype in developing seeds. The mutants had 30–50% less SUS activity and therefore accumulated 40% more sucrose and 50% less fructose at 15 DAF. This did not affect the hexose-P pool, but led to 30–70% less starch in embryo and seed coat. Lipids were 55% higher in both mutants at 9–15 DAF. It seems that sucrolysis via SUS is not required for oil or protein synthesis but rather for channeling carbon toward ADP-glucose and starch in seeds. Metabolite profiling with GC–TOF revealed specific downstream changes in primary metabolism as a consequence of signaling or regulatory fine-tuning. While sucrose increased, hexoses and specific amino acids decreased reciprocally. There was a developmental shift regarding an earlier timing of dry weight accumulation, germinative maturity, oil deposition, sugar levels, transient starch buildup, and protein storage. Nevertheless, final seed size and composition were unaltered due to an earlier cessation of growth, thus giving rise to an apparent silent phenotype of mature mutant seeds. We conclude that SUS is important for metabolite homeostasis and timing of seed development, and propose that an altered sucrose/hexose ratio can modify carbon partitioning and the pattern of storage compounds in Arabidopsis.     

The performance of carrot seeds in relation to their viability

The germination and emergence characteristics, and early seedling growth, of carrot seeds cv. Chantenay red-cored from different sources with a range of germination from 54–94%, was compared. Seeds from protected crops (mean temperature of growth 21°C, r.h. 45–70%) gave higher percentage germination than those from crops grown outdoors (mean temperature 15°C, 70–100% r.h.). Germination was also higher from mature (seed moisture content at harvest 20% or lower) than immature seed (seed moisture content at harvest between 20 and 60%). High percentage germination (>90%) was associated with low mean germination times and low spreads of germination times whilst the reverse was true for low percentage germination. Similar relationships were found for seedling emergence characters in the field although a lower proportion of the viable seeds produced seedlings from slowly than rapidly germinating seed lots. In general, seed lots having a low percentage germination gave greater variability in plant weight than those of higher percentage germination. There was no effect of seed source on radicle or shoot relative growth rates or on post-emergence seedling growth rates.