Desert species adapted for dispersal and germination during floods: Experimental evidence in two Astrophytum species (Cactaceae)

Dispersal of organisms by water is known as hydrochory and is an important means of seed transportation. Here we present experimental evidence of possible dispersal and germination of seeds in water for arid land cacti. Seeds from Astrophytum spp. have unusual traits for plants from arid environments, traits that are typical of water dispersed seeds. Astrophytum capricorne and A. ornatum seeds disperse by floating in the Chihuahuan Desert. Seeds of other native species (Acacia farnesiana, Opuntia streptacantha and Prosopis laevigata (mesquite) were used as control, as well as commercial beans (Phaseolus vulgaris). Seed structures of A. capricorne and A. ornatum are consistent with water dispersal (hat shape, a funicular envelope covering a prominent hilum, and air chambers throughout the tegument). Seeds of A. capricorne showed the greatest water permeability, higher than those of A. ornatum which had similar permeability to common beans and mesquite. Acacia farnesiana and Opuntia streptacantha had the lowest permeability. Seeds of A. capricorne without a funicular layer floated more than those with a funicular layer. All seeds of A. ornatum floated initially, but 10% of those with funicular layer sank after a 30 days floating period. P. laevigata seeds floated also, but less than those of Astrophytum spp., and seeds of A. farnesiana and Phaseolus vulgaris did not float. Germination of A. capricorne was high under all treatments, both when immersed in a water layer or only moistened. Germination of A. ornatum was generally low (<30%), with <5% germination for seeds immersed in water and for those placed on a water film. Seeds of A. capricorne germinated slower in a water layer than in other treatments. No difference in germination rate was found across all treatments in A. ornatum.     

Embryo structure,seed traits,and productivity of relict vine Aristolochia contorta (Aristolochiaceae)

The herbaceous vine Aristolochia contorta (Aristolochiaceae) is a rare plant with a fragmented area in East Asia. Confocal laser-scanning microscopy and stereomicroscopy were employed to examine the seed and embryo structure. This is the first research on embryo anatomy in Aristolochia. Shape and structure of this embryo may be phylogenetically significant. The seed structure of A. contorta is similar to that of other Aristolochia species. Considerable variation is found for productivity traits; the average seed set is rather low (26.7%). The studied seed and embryo traits of A. contorta are found to be variable, but may be potentially informative at the genus and species level. The restricted productivity and reduced population size threaten the continued survival of A. contorta and, perhaps, the butterfly that depends on it.     

Serotiny and seed germination in three threatened species of Mammillaria (Cactaceae)

Individuals of some species of Mammillaria (Cactaceae) store some seeds on the plant over periods exceeding 1 year (serotiny). We examined the phenomenon of serotiny and germination behaviour of three rare and endangered Mammillaria species that occur in central Mexico. The species with the highest seed retention was Mammillaria solisioides, whose individuals kept on average 24% of their total seed crop throughout their observable lifetime. Individuals of Mammillaria napina and Mammillaria hernandezii did not differ in their degree of seed retention (about 5%). In M. solisioides and M. hernandezii, seed germination declined significantly with seed age, whereas in M. napina germination increased slightly. In all three species, over 70% of retained seeds were still alive after 8 years. Increasing fractions of dormant seeds were observed with seed age in M. solisioides and M. hernandezii, whereas in M. napina this fraction followed the opposite trend. All three species showed strict light dependence for germination. Serotiny was positively correlated with the harshness of the environment when species and populations were assumed independent. However, these correlations were not significant at the 5% level when the degree of relatedness of species and populations was taken into account using phylogenetically independent constrasts. We hypothesise that serotiny in these species represents a mechanism by which they can cope with a harsh, unpredictable environment. To our knowledge, this is the first assessment of serotiny in cacti.     

刺五加胚和胚乳发育的研究

刺五加Eleutherococcus senticosus(Rupr .et Maxim．)Maxim的胚胎发生类型为茄型。其卵细胞受精后,合子经历15天左右的休眠期才进行第一次分裂。合子分裂通常发生在胚乳细胞化之后,经棒形胚、球形胚、至果实成熟时发育到心形胚。棒形胚后期至心形胚初期,胚柄最为发达。刺五加的胚乳发育类型为核型。其初生胚乳核的休眠期为1天左右。当胚乳游离核数目增加到200至300时,胚乳以自由生长细胞壁的方式细胞化,胚乳细胞以典型的有丝分裂方式进一步增殖,增加细胞数目。球形胚时期,胚乳细胞内开始贮藏营养物质。少数种子的胚乳里存在巨大细胞核的异型胚乳细胞。在胚乳游离核为32至64个时,分化出珠被绒毡层;球形胚时期,珠被绒毡层解体。珠被绒毡层解体后,胚乳表层细胞分化为分泌层。球形胚至心形胚阶段,约有5％的种子里,胚与胚乳组织发生弥散样降解。成熟果实中,含有大量的瘪粒种子和虫咬种子;饱满种子率为40％左右。饱满种子中,胚乳组织占据种子体积的绝大部分,胚所占比率很小。讨论了不同发育时期胚和胚乳的营养供应。     

Flowering and pollination patterns of Magnolia denudata with emphasis on anatomical changes in ovule and seed development

Magnolia denudata is an excellent ornamental and ecologically important tree that exhibits low fecundity because of seed abortion. To better understand the cause of this low sexual reproduction, pistil receptivity, and pollen quality were investigated using culture medium and histochemical staining, with emphasis on pollen viability at different flowering stages. Cross-pollination, geitonogamy, and self-pollination were undertaken on flowers of sample trees. We found that flowers produce a large number of pollen grains available for effective pollination. Pollen viability varied at different stages, but was highest during stage 3, in which the stamens detach from the axis, the anthers dehisce, and the gynoecium stigma starts to fade. The duration of pistil receptivity was approximately 5 h. These results indicate that poor fertilisation may occur if the optimum period for pollination is missed or the stigma receives poor-quality pollen grains from stages other than stage 3. Although dichogamy avoids selfing, it has led to considerable geitonogamy. Our anatomical analysis of ovule and seed development further revealed that fertilisation occurred in samples with geitonogamy, but that the embryo degenerated in the torpedo stage, demonstrating that aborted seeds were produced by fertilisation, rather than by pseudogamy. Our results provide new insights into the mechanism of reduced seed set under natural conditions.     

Structural aspects of ovule and seed development and nonrandom abortion inMelilotus officinalis (Fabaceae)

Summary Only one ovule matures into a seed inMelilotus officinalis. Although eight ovules form within an ovary, only the basal ovule develops into a mature seed, whereas the other ovules degenerate. The investigation of ovule and seed structure at different developmental stages and a comparison of quantitative characters of differently fated ovules within an ovary were undertaken by light, phase contrast, and fluorescence microscopy. In this species, campylotropous ovules develop simultaneously on marginal placentae in an apocarpous unilocular gynoecium. Megasporo- and megagametogenesis proceed normally and are completed in bud. The maturation of the Polygonum type embryo sac takes place after the flower opens. Shortly before fertilization, synergids show signs of degeneration in all ovules. At this stage, neither the structure nor the sizes of ovules within one ovary differ significantly. In spite of this, only the basal ovule develops into a seed. Rarely, one of the upper-situated ovules or the basal and another ovule mature into seeds. Seed enlargement is insignificant until the stage when globular embryo and nuclear endosperm are formed. At the seed-filling stage, other ovules have collapsed and the seed gradually comes to occupy the total volume of the pod. The fruit-to-seed length ratio decreases considerably during seed ripening. At fertilization, ovary length is four times greater than ovule length. In the mature state, the fruit and seed lengths are approximately equal. Seed size and weight diminish with an increase in seed number within a pod, although pod size remains constant. It is assumed that nonrandom abortion of young seeds inM. officinalis is under maternal control and is not related to structural abnormalities in ovule development or with limitation in pollen. We suppose that evolution of this species may have proceeded in the direction of a decrease in seed number and an increase in its sizes, which may play an important role in seed dispersal and seedling establishment.     

Antigenic patterns of seed proteins in Opuntioideae (Cactaceae)

The antigenic patterns recognised by Western blotting in seed proteins of species of Opuntioideae (Cactaceae) were analysed in an attempt to evaluate their usefulness in systematics. Total protein profiles were also analysed by SDS–PAGE. The resulting similarity and distance matrices were further used to carry out Cluster Analysis (UPGMA) and Principal Coordinates Analysis. Populations of Opuntia cardiosperma were found to exhibit a prominent morphological uniformity, a unique electrophoretic pattern and a uniform antigenic pattern. The latter was obtained using anti-O. cardiosperma as antiserum. Results from the qualitative and quantitative interspecific analyses of antigenic profiles helped to characterise all the species studied. Tephrocactus articulatus and Cylindropuntia imbricata evidenced lower affinity with O. cardiosperma than the species of Opuntia s.s. Our results demonstrate that in Cactaceae, Western blotting analysis broadens the usefulness range of immunological techniques at the specific level and complements the information collected from electrophoretic profiles.     

Ovule and seed development of Lacandonia schismatica (Lacandoniaceae)

Ovule and seed development is described for Lacandonia schismatica, a species whose androecium is surrounded by the gynoecium. The ovule in each carpel is basal, anatropous, tenuinucellate, and bitegmic. The female gametophyte is formed by the micropylar megaspore cell, after a coenocytic stage of the four megaspore nuclei. The mature female gametophyte has the normal complement of seven cells and eight nuclei. We propose a new type of female gametophyte development on the basis of the coenocytic stage of the tetrad, the cellularization of the tetrad, and the survival of the micropylar spore. At seed dispersal time, the embryo has ~10-20 cells. Endosperm development is of the nuclear type. At maturity, endosperm cells show starch and protein inclusions as well as polysaccharides in their thick walls. The seed coat is formed from the outer integument; the inner one disappears. The exotesta contains tannin. The fruit (achene) wall is two-layered. The maturation of the fruits in a flower is synchronous, and they separate from the receptacle for dispersal.     

Aerial seed bank and dispersal traits in Anthemis chrysantha (Asteraceae), a critically endangered species

Mature plant density and fruit production were monitored in the main population of four successive cohorts of the endangered winter annual Anthemis chrysantha (Asteraceae) in southeastern Spain. Experiments were conducted with artificial rainfall and a wind tunnel to determine the temporal and spatial dispersal pattern of the species and the relationship with rain and wind.     

Paternal regulation of seed development in wheat hybrids

Summary Diallel crosses among Triticum boeoticum (4 lines from different geographical areas), T.urartu, Aegilops squarrosa and Ae. speltoides exhibited reciprocal differences in hybrid seed morphology, endosperm development, and embryo viability. T. urartu and Ae. squarrosa as females with T. boeotiaum and Ae. speltoides lead to shrivelled inviable seed. T.boeoticum accessions as female with Ae.speltoides also lead to shrivelled seeds. The reciprocal crosses produced plump seeds which either resembled the maternal parent or showed size differences. By altering the endospermic genome ratios, hybrid seeds with 1 (PF)/1 (PM) showed extreme shrivelling whereas those with 4 (PF)/1 (PM) were medium shrivelled to plump. Genetic experiments involving hybrids of T. boeoticum, T. urartu and T. monococcum showed that a factor is present in pollen or male gametes, which shows dosage effect and which, by interacting with the maternal genome, leads to endosperm abortion.     

Degree of pectin methyl esterification in endosperm cell walls is involved in embryo bending in Arabidopsis thaliana

The endosperm is a transitory structure involved in proper embryo elongation. The cell walls of mature seed endosperm are generally composed of a uniform distribution of cellulose, unesterified homogalacturonans, and arabinans. Recent studies suggest that changes in cell wall properties during endosperm development could be related to embryo growth. The degree of methyl esterification of homogalacturonans may be involved in this endosperm tissue remodelling. The relevance of the degree of homogalacturonan methyl esterification during seed development was determined by immunohistochemical analyses using a panel of probes with specificity for homogalaturonans with different degrees of methyl esterification. Low-esterified and un-esterified homogalacturonans were abundant in endosperm cells during embryo bending and were also detected in mature embryos. BIDXII (BDX) could be involved in seed development, because bdx-1 mutants had misshapen embryos. The methyl esterification pattern described for WT seeds was different during bdx-1 seed development; un-esterified homogalacturonans were scarcely present in the cell walls of endosperm in bending embryos and mature seeds. Our results suggested that the degree of methyl esterification of homogalacturonans in the endosperm cell wall may be involved in proper embryo development.     

AtNG1 encodes a protein that is required for seed germination

The pentatricopeptide repeat (PPR) family of eukaryotic proteins has numerous members in plants and is important for plant development. In the present study, we cloned a novel PPR gene, designated AtNG1, and characterized the ng1 Arabidopsis mutant. Morphological and structural observation of an ng1 mutant revealed that its sexual reproduction and seed formation processes are essentially normal. The mature embryonic root of ng1 is fully developed and has a well-differentiated structure; however, ng1 seeds cannot germinate, even when supplied with supplemental hormones and nutrition. Further investigation showed that embryo expansion and root cell elongation fails to occur after water imbibitions. Transient gene expression analysis indicated that AtNG1 localizes in mitochondrion. This implies that the deficiency of mitochondrion function might be the reason for the failed seed germination. Thus, our finding confirmed that AtNG1 plays a critical role in the early process of seed germination.     

Water limitation effect on seed development and germination in Trigonella coerulea (Fabaceae)

The annual garden spice legume Trigonella coerulea was subjected to controlled drought conditions to investigate the influence of resource limitation on flowering, seed production and germination. Limitations in water availability significantly reduced plant height and the number of fruits produced. However, treatments had no significant effect on seed set within the fruit. Fruit number per plant, rather than seed number per fruit was affected by resource limitation. Plants growing under water deficiency had higher flower abortion rates. Simultaneously, the size and germination rate of the seeds were decreased. In terms of reproductive success T. coerulea was not able to adapt reproduction to water shortage. Increase in moisture had no significant effect on seed quantity and quality when compared to the control.     

Developmental anatomy and morphology of the ovule and seed of heliconia (heliconiaceae, zingiberales)

The developmental anatomy and morphology of the ovule and seed in several species of Heliconia were investigated as part of an embryological study of the Heliconiaceae and to provide a better understanding of their relationships with the other families of the Zingiberales. Heliconia species have an ovule primordium with an outer integument of both dermal and subdermal origin. The archesporial cell is divided into a megasporocyte and a single parietal cell, which in turn are divided only anticlinally to form a single parietal layer, disintegrating later during gametogenesis. The embryo sac was fully developed prior to anthesis. In the developing seed, the endosperm was nuclear, with wall formation in the globular stage; a nucellar pad was observed during embryo development, but later became compressed. The ripe fruit contained seeds enveloped by a lignified endocarp that formed the pyrenes, with each pyrene having an operculum at the basal end; the embryo was considered to be differentiated. Most of these characteristics are shared with other Zingiberales, although the derivation of the operculum from the funicle and the formation of the main mechanical layer by the endocarp are unique to the Heliconiaceae.     

Ecological and morphological seed traits of Polygala sardoa and P. sinisica: A comparative study on two endemic species of Sardinia

Polygala sardoa Chodat and P. sinisica Arrigoni (Polygalaceae) are two exclusive endemics to Sardinia and P. sinisica is affiliated under “Critically Endangered” in the IUCN Red Lists. In this work comparative studies on two populations of P. sardoa and in the only one of P. sinisica were carried out. In particular, seed output calculations and morphometric measurements on seed and elaiosome sizes were carried out. The effect of chipping, constant (15 °C) and alternating (25/10 °C) temperature regimes and light (8 h irradiance/day and dark) on germination were investigated. Seedling vigor assessments and a study on seed persistence in the soil were also carried out. P. sinisica had a higher seed output than P. sardoa and P. sinisica seeds were smaller than those of P. sardoa, with lower elaiosome area and elaiosome ratio values. Neither species had alternating temperature or light requirements for germination. P. sardoa achieved high germination percentages (> 80%) at all the tested conditions. P. sinisica had a lower maximum germination (<60%), suggesting the presence of physiological dormancy, and took more than twice as long to reach the maximum germination percentage. Seedlings of P. sardoa were larger than those of P. sinisica, and neither species formed a persistent soil seed bank. These new data may help implement effective conservation measures for these two species and, more generally, for threatened endemic species of this genus.     

Ovule,fruit and seed development in Abolboda (Xyridaceae,Poales): implications for taxonomy and phylogeny

Xyridaceae belongs to the xyrid clade of Poales, but the phylogenetic position of the xyrid families is only weakly supported. Xyridaceae is divided into two subfamilies and five genera, the relationships of which remain unclear. The development of the ovule, fruit and seed of Abolboda spp. was studied to identify characteristics of taxonomic and phylogenetic value. All of the studied species share anatropous, tenuinucellate and bitegmic ovules with a micropyle formed by the inner and outer integuments, megagametophyte development of the Polygonum type, seeds with a tanniferous hypostase, a helobial and starchy endosperm and an undifferentiated embryo, seed coat derived from both integuments with a tanniferous tegmen and a micropylar operculum, and fruits with a parenchymatous endocarp and mesocarp and a sclerenchymatous exocarp. Most of the ovule and seed characteristics described for Abolboda are also present in Xyris and may represent a pattern for the family. Abolboda is distinguished by the ovule type, endosperm formation and the number of layers in the seed coat, in agreement with its classification in Abolbodoideae. The following characteristics link Xyridaceae to Eriocaulaceae and Mayacaceae, supporting the xyrid clade: tenuinucellate, bitegmic ovules; seeds with a tanniferous hypostase, a starchy endosperm and an undifferentiated embryo; and a seed coat with a tanniferous tegmen. A micropylar operculum in the seeds of Abolboda is described for the first time here and may represent a synapomorphy for the xyrids. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 144–154.     

Seed germination in Marathrum schiedeanum and M. rubrum (Podostemaceae)

Marathrum schiedeanum and Marathrum rubrum are annual Podostemaceae, thus their seeds are important to their dispersal and persistence in their habitat. We assessed the effect on germination of (1) light (white, red and far red) and darkness, (2) temperature (15, 20, 25, 30 °C and alternating 20/30 °C), (3) osmotic potential (0 to −0.8 MPa), (4) proximity to moisture sources and (5) seed storage. Seeds of M. schiedeanum and M. rubrum were non-dormant and had a high germination capacity (96%). Seeds were positive photoblastic; at 15 °C germination drop to zero, and germination rate was slower at 20 °C and at 20/30 °C than at 25 °C. A small proportion of seeds of both species germinated even at osmotic potentials as low as −0.6 MPa (11%) for M. rubrum and −0.8 MPa (70%) for M. schiedeanum. Seeds germinated only when near to the source of moisture (91.3–87.1% and 53.3–35.6% for M. schiedeanum and M. rubrum, respectively) and 2 years in dry storage did not modify their capacity to germinate. At the beginning of the rainy season, light and temperature in the rivers may be high enough for germination. The ability to germinate at low osmotic potential may be related to early germination during the rainy season. This may be because the seed mucilage assists in diffusion of water from the substrate to the seed. Both species germinated faster at −0.06 MPa, than in distilled water, which may indicate appropriate conditions for germination of these short-lived species.     

Systematic implication of seed micromorphology in Arenaria (Caryophyllaceae) and allied genera

Variations in seed micromorphology of 64 taxa of Arenaria and allied genera were examined using scanning electron microscopy (SEM) in order to evaluate their diagnostic value for systematic studies in the genus and providing additional evidence on delimitation of natural groups. Significant diversity was found in seed coat morphology both among and within genera. Size, color, and shape of seeds appear to be of low importance at species level but provide useful evidence at generic rank. Cerastium and Stellaria are well distinguished from Arenaria by serrate seed margins and solid columellae on testa cells. The sculpturing pattern renders most characters potentially applicable in taxonomy and phylogeny assessment in Arenaria. The studied taxa show four basic types of sculpturing, viz. colliculate, rugose, papillate, and smooth. The colliculate type which is characteristic for the core group of Arenaria (A. subg. Arenaria) can be further divided into five subtypes. The smooth seed surface with indistinct cell boundaries represents a synapomorphy for A. subg. Leiosperma, whereas the winged seeds along with rugose surface characterize subg. Odontostemma. Otherwise seed micromorphology rarely provides adequate evidence for separation of natural groups, especially at sectional rank. We hypothesize that the seeds are highly polymorphic in certain groups of taxa showing recent adaptive radiation, such as the annual species of Arenaria, and do not provide strong support for natural groups within the genus.     

Abortive seed development in Ulmus minor (Ulmaceae)

Three genotypes of field elm ( Ulmus minor ) were studied to determine the structural basis of seed abortion in this species. In the non-abortive control, P-VV1, the pattern of seed development is similar to many flowering plants. The embryo progresses through defined morphological stages leading to developmental arrest as the seed matures. Storage products are abundant within embryo cells. Endosperm development is similar to the nuclear type; however, a more extensive cellularization of the endosperm occurs prior to it being crushed by the expanding embryo. For the abortive genotypes, M-SF1 and V-JR1, abnormalities in endosperm development are found. This is judged by the early cellularization and the massive synthesis of the PAS-positive material in the cellular endosperm. In these abortive genotypes, embryo development is delayed and storage products failed to accumulate within embryo cells. After seed desiccation, no living embryo tissue remains within the seed coat in the abortive genotypes.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 455–467.