Seed micro‐morphology and its systematic significance in Gypsophila (Caryophyllaceae) and allied genera

In order to examine the systematic application of seed‐coat micro‐morphology in Gypsophila and allied genera, seed surfaces of 30 species and two varieties representing four genera of tribe Caryophylleae were examined with scanning electron microscope. The seeds of examined species range between 0.5–2.1 mm in length and 0.3–2.4 mm in diameter. The exomorphology of the seed coat shows two distinctive cell patterns. The epidermis is constructed either of elongated polygonal or of broad polygonal cells. The elongated type is the most common among the studied species, but the variation in alignment of testa cells, their size and shape as well as the density of protuberances may provide further information and useful diagnostic characters at generic and specific rank. The testa cells in Gypsophila and Saponaria are shallowly undulate, deeply undulate, lobed and armed at anticlinal walls. Deeply undulate anticlinal walls were observed in both Gypsophila and Ankyropetalum and a few species of Saponaria. Non‐ or indistinctly grooved anticlinal walls is the more common type in Allochrusa. Seed‐coat characters support the separation of Gypsophila and Saponaria to some extent but disagree with recognition of Ankyropetalum as a genus separate from Gypsophila.     

Mikromorphologie der Samenoberflächen der GattungEuphorbia

Seed coat surfaces of 127 species, representing 23 sections of the genusEuphorbia L. (Euphorbiaceae) have been examined with the scanning electron microscope (SEM). Various surface patterns and cell wall structures are described. In some groups the seed coat is rather uniform (e.g. in sect.Euphorbium), in other sections even closely related species can be separated by seed coat morphology (e.g.Tithymalus). The taxonomic application of testa micromorphology and the possible systematic position of certain taxa are briefly discussed. The seed coat cells of all species in sect.Tithymalus show remarkable intercellulares filled with small particles, which are described in detail for the first time.

     

SEED COAT MORPHOLOGY IN CORDYLANTHUS (SCROPHULARIACEAE) AND ITS TAXONOMIC SIGNIFICANCE

A study of seed coat sculpturing in Cordylanthus (Scrophulariaceae—Rhinantheae) using the scanning electron microscope (SEM) shows that seed surface patterns are characteristic and constant for a given species or a group of species. Seeds of 23 species were examined and classified into four types (irregularly crested, deeply reticulate, shallowly reticulate, and irregularly striate) based on differences in the reticulated seed coat. Anatomical studies of sections with the light microscope show that the reticular patterns result from enlargement of the epidermal cells of the integument, followed by formation of characteristic wall patterns through lignification. Seed characters furnish useful data for formulating the taxonomy of Cordylanthus both on the sectional and subsectional level and for the delimitation of certain species. Seed coat morphology offers evidence for including the genus Dicranostegia in Cordylanthus and for maintaining C. hispidus and C. palmatus as separate species. The distinctness of two seed coat types within section Cordylanthus suggests two major phylogenetic lines within this section.     

勾儿茶属和小勾儿茶属(鼠李科)的果实及种子形态研究及其系统学意义

在扫描电镜和解剖镜下研究鼠李科(Rhamnaceae)勾儿茶属(Berchemia)和小勾儿茶属(Berchemiella)共17种植物的果实和种子形态.结果表明：小勾儿茶属的核果1室,具1枚种子,勾儿茶属大多数种的核果2室,每室具1枚种子,但多叶勾儿茶(B．polyphylla)的核果1室较大,具1枚种子,另一室较小而没有种子,该种可能是连接勾儿茶属和小勾儿茶属的中间类群.这两个属的种子形状通常为不规则长椭圆形,种皮纹饰可划分为光滑或几乎光滑、具不明显或稀疏的条纹以及具明显的条纹或沟这三种类型.种皮纹饰的差异对这两个属属下种类的划分具有一定的意义.讨论了与前人研究结果不同之处和可能的原因.     

Micromorphology of fruits and seeds of Iranian Geranium (Geraniaceae), and its systematic significance

The fruit and seed micromorphology of 22 species of Geranium, representing the eight sections of the genus represented in Iran (G. sectt. Dissecta, Geranium, and Tuberosa of subgen. Geranium; sectt. Batrachioidea, Divaricata, Lucida, Ruberta and Trilopha of subgen. Robertium), have been examined by scanning electron microscopy (SEM). Macro‐ and micromorphological characters, including fruit and seed shape, size, color, hair type and density, mericarp ornamentations, hilum position, seed coat pattern, epidermal cell shape, and anticlinal and periclinal cell walls, are presented. Two microsculpturing patterns are recognized on the mericarp surface: reticulate and pusticulate. The micromorphology of the seed coat showed four distinctive cell patterns. The seed epidermis is constructed either of polygonal, elongated polygonal, or square to rectangular cells. The polygonal type is the most common among the studied species, but the variation in testa cell characters, their size and shape, may provide further information and useful diagnostic characters at specific and infraspecific rank. The shape and color of the seeds are, however, of little systematic value. Fruit characters were found to be important for separating taxa at infrageneric rank and our results show that the species can be separated into subgenera and sections based on fruit morphology.     

Rodent seed predation promotes differential recruitment among bird-dispersed trees in temperate secondary forests

We investigated the role of seed predation by rodents in the recruitment of the fleshy-fruited trees Taxus baccata, Ilex aquifolium and Crataegus monogyna in temperate secondary forests in NW Spain. We measured the densities of dispersed seeds, early emerged seedlings, established recruits and adults, at four sites over a period of 2 years. Seed predation among species was compared by seed removal experiments and analysis of rodent larder-hoards. The three species differed markedly in local regeneration patterns. The rank order in the seed rain following decreasing seed density was Ilex, Taxus and Crataegus. However, Crataegus established 3.3 times more seedlings than Taxus. For all species, there was a positive linear relationship between the density of emerged seedlings and seed density, suggesting that recruitment was seed- rather than microsite-limited. A consistent pattern of seed selection among species was exerted by rodents, which preferred Taxus and, secondarily, Ilex seeds to Crataegus seeds. Predation ranking was the inverse of that of seed protection against predators, measured as the mass of woody coat per mass unit of the edible fraction. Recruitment potential, evaluated as the ratio of seedlings to seeds, was negatively related to seed predation, with the rank order Crataegus > Ilex > Taxus. The selective early recruitment limitation exerted by predation may have a demographic effect in the long term, as judged by the positive relationship between early seedling emergence and the density of established recruits. By modulating the pre-emptive competition for seed safe sites, rodents may preclude the progressive exclusion of species that produce low numbers of seeds (i.e. Crataegus) by those dominant in seed number (i.e. Ilex, Taxus), or at least foster the evenness for site occupation among seedlings of different species.     

Seed coat micromorphology of South American species of Hybanthus (Violaceae)

The seed‐coat of several species of Hybanthus subg. Ionidium from South America were studied by SEM (scanning electronic microscopy). Three different levels of sculpture were observed on the seed coat. The micro‐morphological patterns showed great variation between the species and should be considered as additional characters in future taxonomic treatments. The presence of a conspicuous elaiosome was observed; this structure is well known in the genus Viola and is most likely related to seed dispersal mediated by ants.     

Comparative seed morphology and character evolution in the genus Lysimachia (Myrsinaceae) and related taxa

Summary We investigated seed morphology in 34 species of the genus Lysimachia and in 14 species and two subspecies of six additional genera (Anagallis, Ardisiandra, Asterolinon, Glaux, Pelletiera, Trientalis), which have been shown to be closely related to, or are placed within Lysimachia in previous molecular studies. We studied seed shape, seed coat structure, and seed coat surface patterns. Three major types of seed shape were identified: (1) sectoroid, (2) polyhedral, and (3) coarsely rugose with concave hilar area. In addition, seeds may be keeled or winged. The outer layer of the seed coat is either sponge-like and adhering only loosely to the inner seed coat or it is thin and tightly adhering to the underlying tissue. Seed surface patterns can be divided into six main types: (1) reticulate, (2) tuberculate, (3) vesiculose, (4) colliculate, (5) undulate, or (6) poroid-alveolate. Seed surface patterns are mostly congruent with molecular phylogenetic relationships. A reticulate surface pattern is diagnostic of, e.g. Lysimachia subgenera Palladia and Hawaiian Lysimachiopsis. Mapping seed characters onto a recent phylogenetic tree, reveals that they provide potentially synapomorphic character states for various subclades of Lysimachia. Salient examples include a rugose seed shape, which turns out to be synapomorphic for the clade comprising the genus Pelletiera plus Asterolinon linum-stellatum and a sponge-like outer seed coat layer, which characterizes a clade with Lysimachia vulgaris, L. thyrsiflora, and L. terrestris, with an analogue that apparently evolved in parallel in Trientalis europaea. We also discuss possible habitat factors that may have favored the independent evolution of particular seed types such as winged seeds in various lineages.     

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     

Does fire drive fatty acid composition in seed coats of physically dormant species?

• Seed dormancy is the key driver regulating seed germination, hence is fundamental to the seedling recruitment life-history stage and population persistence. However, despite the importance of physical dormancy (PY) in timing post-fire germination, the mechanism driving dormancy-break within seed coats remains surprisingly unclear. We suggest that seed coat chemistry may play an important role in controlling dormancy in species with PY. In particular, seed coat fatty acids (FAs) are hydrophobic, and have melting points within the range of seed dormancy-breaking temperatures. Furthermore, melting points of saturated FAs increase with increasing carbon chain length. We investigated whether fire could influence seed coat FA profiles and discuss their potential influence on dormancy mechanisms.
• Seed coat FAs of 25 species within the Faboideae, from fire-prone and fire-free ecosystems, were identified and quantified through GC–MS. Fatty acid profiles were interpreted in the context of species habitat and interspecific variation.
• Fatty acid compositions were distinct between species from fire-prone and fire-free habitats. Fire-prone species tended to have longer saturated FA chains, a lower ratio of saturated to unsaturated FA, and a slightly higher relative amount of FAs compared to fire-free species.
• The specific FA composition of seed coats of fire-prone species indicated a potential role of FAs in dormancy mechanisms. Overall, the distinct FA composition between fire-prone and fire-free species suggests that chemistry of the seed coat may be under selection pressure in fire-prone ecosystems.

     

Factors Affecting Attachment of Enterobacter cloacae to Germinating Cotton Seed

Abstract Attachment of Enterobacter cloacae EcCT-50,—a biological seed protectant used to control the seed-rotting fungi, Pythium ultimum—to cotton seed was examined using conventional fluorescent microscopy (CFM), scanning electron microscopy (SEM), and laser scanning microscopy (LSM). In sand microcosms, E. cloacae quickly attached to the seed coat, with maximum attachment, 3 to 5 h after inoculation at 24°C. In contrast, initial attachment of non-bacterized seed by Pythium ultimum was not observed until 6 h (and not until 8 h on bacterized seeds). Comparison of the movement of E. cloacae and P. ultimum in seed exudate gradient semi-soft agar showed faster movement by the bacterium within the first 6 h, and reduction of P. ultimum hyphal and germ tube growth in the presence of the bacterium. Microscopic observation of the seed coat revealed an early, intimate association, mediated, in part, by fimbriae, and confirmed a loose association of E. cloacae with the seed coat previously reported. Spatially, the attached E. cloacae cells were distributed over the entire surface of the seed coat, but were especially abundant in the groves and near cracks where water imbibition and seed exudate release may occur. As the seed germinated and exposed various seed tissues, the bacterium rapidly attached to these tissues. Attachment of the bacterium to the surface of intact germinating seeds, excised seed coat, polystyrene, and glass was 300, 110, 51, and <1 cell field⁻¹ 3 h⁻¹, respectively, suggesting that attachment is enhanced by seed germination. Attachment of E. cloacae to the seed coat was optimum in sands with high water concentrations, at temperatures of 18 to 30°C, and at times that corresponded with optimum water imbibition during germination. Using several assays, attachment was shown to be enhanced by seed exudate, and compounds such as methanol, fructose, and calcium. The results suggest that the release of certain nutrients and water imbibition during germination may play a role in the rapid attachment to the seed by E. cloacae. The ability of E. cloacae to rapidly move and attach to the seed coat may be related to its ability to function as a biocontrol agent. Received: 24 April 1997; Accepted 29 October 1997     

Seed reserve composition in 19 tree species of a tropical deciduous forest in Mexico and its relationship to seed germination and seedling growth

Background and Aims

The size and composition of seed reserves may reflect the ecological strategy and evolutionary history of a species and also temporal variation in resource availability. The seed mass and composition of seed reserves of 19 co-existing tree species were studied, and we examined how they varied among species in relation to germination and seedling growth rates, as well as between two years with contrasting precipitation (652 and 384 mm).

Methods

Seeds were collected from a tropical deciduous forest in the northwest of Mexico (Chamela Biological Station). The seed dry mass, with and without the seed coat, and the concentrations of lipids, nitrogen and non-structural carbohydrates for the seed minus seed coat were determined. The anatomical localization of these reserves was examined using histochemical analysis. The germination capacity, rate and lag time were determined. The correlations among these variables, and their relationship to previously reported seedling relative growth rates, were evaluated with and without phylogenetic consideration.

Key Results

There were interannual differences in seed mass and reserve composition. Seed was significantly heavier after the drier year in five species. Nitrogen concentration was positively correlated with seed coat fraction, and was significantly higher after the drier year in 12 species. The rate and lag time of germination were negatively correlated with each other. These trait correlations were also supported for phylogenetic independent contrasts. Principal component analysis supported these correlations, and indicated a negative association of seedling relative growth rate with seed size, and a positive association of germination rate with nitrogen and lipid concentrations.

Conclusions

Nitrogen concentration tended to be higher after the drier year and, while interannual variations in seed size and reserve composition were not sufficient to affect interspecific correlations among seed and seedling traits, some of the reserves were related to germination variables and seedling relative growth rate.     

Allium aetnense (Amaryllidaceae), a new species from Sicily

A new species from Mt Etna (Sicily), Allium aetenense of A. sect. Codonoprasum, is described and illustrated. It is a diploid species (2n = 16) that grows on basaltic rocks in the mountain belt and shows close relationships with A. tenuiflorum of the Italian peninsula. Its morphology, karyology, leaf anatomy, seed coat microsculpturing, ecology and taxonomic position are examined.     

Variation of proline rich cell wall proteins in soybean lines with anthocyanin mutations

The I locus controls inhibition of anthocyanin accumulation in the epidermal cells of the soybean seed coat and affects abundance of PRP1, a proline-rich cell wall protein in the seed coat. Saline-soluble PRP1 is abundant in the developing seed coats of cultivar Richland (homozygous I, yellow), while it is significantly decreased in the pigmented isogenic mutant T157 (homozygous i, imperfect black). In this report, we examined soluble PRP1 in several cultivars containing alleles of the I locus which affect spatial distribution of pigmentation in the seed coat. We also characterized PRP1 in isolines with allelic variants of several other loci involved in seed coat pigmentation, including T and Im. The T gene is pleiotropic and affects both pubescence color and seed coat pigmentation and structure. Soluble PRP1 was abundant in the developing seed coats of lines with yellow seed (I or i ⁱ alleles) regardless of pubescence color, just as in Richland. Likewise, soluble PRP1 was decreased in pigmented seed coats (i ^k or i alleles) with grey (t) pubescence, as in T157. However, the total seed coat proteins were not extractable from pigmented seed coats with tawny pubescence (i, T genotypes) because they have proanthocyanidins that exhibit tannin properties. The dominant Im allele inhibits seed coat mottling (irregular patches of pigmentation) that occurs if plants are infected with soybean mosaic virus. PRP1 was 35 kDa in mottled (im) isolines and 34 kDa in non-mottled (Im) isolines. PRP2, which is expressed later in seed coat development and in the hypocotyl hooks of soybean seedlings, was also smaller in Im isolines. In summary, some of the anthocyanin mutations affect the quantity of soluble PRP1 polypeptides, while others correlate with structural changes in developmentally regulated proline-rich proteins.     

Novel seed coat lignins in the Cactaceae: structure,distribution and implications for the evolution of lignin diversity

We have recently described a hitherto unsuspected catechyl lignin polymer (C‐lignin) in the seed coats of Vanilla orchid and in cacti of one genus, Melocactus (Chen et al., Proc. Natl. Acad. Sci. USA. 2012, 109, 1772‐1777.). We have now determined the lignin types in the seed coats of 130 different cactus species. Lignin in the vegetative tissues of cacti is of the normal guaiacyl/syringyl (G/S) type, but members of most genera within the subfamily Cactoidae possess seed coat lignin of the novel C‐type only, which we show is a homopolymer formed by endwise β–O–4‐coupling of caffeyl alcohol monomers onto the growing polymer resulting in benzodioxane units. However, the species examined within the genera Coryphantha, Cumarinia, Escobaria and Mammillaria (Cactoideae) mostly had normal G/S lignin in their seeds, as did all six species in the subfamily Opuntioidae that were examined. Seed coat lignin composition is still evolving in the Cactaceae, as seeds of one Mammillaria species (M. lasiacantha) possess only C‐lignin, three Escobaria species (E. dasyacantha, E. lloydii and E. zilziana) contain an unusual lignin composed of 5‐hydroxyguaiacyl units, the first report of such a polymer that occurs naturally in plants, and seeds of some species contain no lignin at all. We discuss the implications of these findings for the mechanisms that underlie the biosynthesis of these newly discovered lignin types.     

Seed coat morphology and evolution in Celtidaceae and Ulmaceae (Urticales)

The seed coat surface morphology of Celtidaceae and Ulmaceae (Urticales) indicates a significant evolutionary diversity.Celtis, Chaetachme andPteroceltis (Celtidaceae) have a unique sculpturing with many crateriform holes; such holes occasionally sparsely occur in seeds ofAphananthe, Gironniera (Celtidaceae) andPlanera (Ulmaceae), but not in those of the nine remaining genera of the two families. The perforated seed coat further occurs in at least some genera of all other urticalean families. A pattern of its occurrence in families and genera suggest that the perforation represents a common archaic feature of all Urticales, rather than a feature derived many times independently within the order. The seed coat of Celtidaceae and Ulmaceae seems to have lately lost the holes probably by a neotenic evolution: one or more times within Celtidaceae, and one time in an ancestral line leading to all Ulmaceae. The derived reticulate seed coat surface sculpturing, which is shared byGironniera (Celtidaceae) and some Ulmaceae, is probably the result of parallel evolution. On the basis of evidence from seed coat morphology and other sources, close relationships ofLozanella, Parasponia andTrema within Celtidaceae, as well as variously distinct positions ofAmpelocera, Aphananthe andGironniera, are also discussed.     

Parasitoid attack of the seed-feeding beetle Bruchus loti enhances the germination success of Lathyrus japonicus seeds

Hard seeds of some legume species can germinate after seed-feeding insects bore through the seed coat and consequently break seed dormancy. Larvae of bruchine beetles are the main seed feeders attacking many legume species. Boring of the hard seed coat by bruchine beetle larvae enhances the germination percentage of legume species, but consuming too much of a single seed may reduce the chances the seed will survive. We hypothesise that the early mortality of bruchine larvae due to parasitism contributes positively to seed germination because larvae are killed before consuming too large a quantity of the seed. Here, we tested this hypothesis using Lathyrus japonicus seeds and Bruchus loti, the main seed feeder attacking this plant. B. loti larvae were mainly parasitised by two species of idiobiont parasitoids—Pteromalus sp. and Dinarmus sp. The seeds from which Pteromalus wasps emerged germinated more successfully than did the seeds from which B. loti adults emerged. B. loti larvae parasitised by the two wasp species consumed the seeds less intensively than did unparasitised larvae. Thus, the results of experiments supported our hypothesis. However, the germination success varied significantly between the seeds from which Pteromalus and Dinarmus wasps emerged. The difference in the size of seeds the two wasp species chose for parasitism may have influenced the germination percentage.     

Ecological implications of seed characteristics of the native Prosopis cineraria and the alien P. juliflora

Seed weight, percentage germination, seedling growth, and nutrient concentrations (Mg, Na, K, Zn, Cu and P) of whole seeds, and of seed coats and embryos separately of two tree species, the native Prosopis cineraria and the invasive alien P. juliflora from semi-arid and arid areas of north and north-west India, were analysed to understand the differences in their ecology. Seeds of P. cineraria were heavier than those of P. juliflora. Percent germination was similar in the two species, but seedling growth was faster in P. juliflora than in P. cineraria. Nutrient concentrations of seeds of the two species were similar (except Cu). Nutrient concentrations in the embryo were higher in P. cineraria, while those in the seed coat were higher in P. juliflora. The relative allocation of nutrients to seed coat was higher in P. juliflora than in P. cineraria. Nutrient-rich embryos and slow growth, along with a staggered seed germination pattern in the native P. cineraria could be linked to delayed establishment as well, in the substratum. Faster growth of the nutrient-poor embryos in P. juliflora along with its simultaneous seed germination pattern, and creation of a favourable microenvironment through leaching of nutrients from a nutrient-rich seed coat can facilitate immediate and successful establishment of this alien species in the invaded habitats.     

Eriocaulon vandaanamense sp. nov. (Eriocaulaceae) from Kerala,India

Eriocaulon vandaanamense (Eriocaulaceae), a new species from a marshy coastal area in the Alappuzha District, Kerala, India, is described and illustrated. The new species is similar to E. truncatum and resembles it in having vertically elongated seed coat cells, but differs mainly in having pale black hoary floral bracts, 3‐lobed hoary male sepals, linear‐falcate, non‐conduplicate hoary female sepals, and small, smooth, pale brown seeds without seed coat appendages.