Genetic diversity in West African okra (Abelmoschus caillei) (A. Chev.) Stevels — Multivariate analysis of morphological and agronomic characteristics

Summary Factor, Principal component and canonical analyses were used to study the extent of genetic diversity among 30 accessions of West African okra. Genetic variability among the accessions proved to be large. Pigmentation of various parts of the accessions and fruit characteristics contributed significantly to the total variation observed. Factor analysis and principal component analysis produced similar results which were substantially different from those produced by canonical analysis. The first three canonical variables accounted for 100% of the total variance while the number of pods per plant, and pod weight primarily accounted for the first canonical variable. Whereas the second canonical variable was primarily loaded by number of seeds per pod and fruit colour, the third canonical variable was comprised of a weight of 100 seeds and number of epicalyx segments.The level of variability observed supports the opinion (Stevels, 1988, 1990) that this okra type constitutes a separate species.     

Agromorphological genetic diversity of Spanish traditional melons

The variability of 62 Spanish landraces (and two hybrids used as reference) was described by analysing 39 morphological traits and eight SSR makers. Results showed that 81% of the examined genetic pool belonged to the inodorus type. Spanish traditional melons presented fruits from flattened and globular shapes to elliptical. Rind colour varied from pale green, almost white and yellow to dark green and almost black. Rind texture varied from smooth to intensely wrinkled. Spanish landraces also had larger fruits (average fruit weight ~2.6 kg) and longer vegetative cycles (117 days to maturity) compared to landraces from other geographical origins. Farmers seem to have focused on selection towards large fruits, which usually requires the longest production cycles. Fruit colour, size and shape seemed to have been determinant in varietal selection. Hierarchical clustering resulted in two main groups (climacteric and non-climacteric). The largest group was composed of 60 accessions of non-climacteric types, which includes the most demanded by national markets, ‘Piel de Sapo’, which fruits were characterized by an ovate or elliptical shape, a green rind, big spots and stains distributed over the whole fruit, a rounded blossom end shape and a very pointed stem end shape. The study demonstrates that the Spanish genetic pool is much more diverse. Wide variability was found in a geographical area with vast historical importance in melon farming. These evaluations has allowed the identification of several uniform groups of non-climacteric cultivars (‘Piel de Sapo’, ‘Mochuelo’, ‘Tendral’, Yellow/White, Winter and Black groups) and a set of highly variable climacteric ones. However, many accessions with singular properties remain unclassified, demonstrating the morphological variability of the studied collection. Melons in Spain have wide variability together with a vast historical importance on farms. Some fruit types, or at least some morphological characters reflected by painters during centuries, have reached the present. The present study confirmed the need to preserve these irreplaceable genetic resources and continue their study and evaluation for valuable traits which could enhance farmer’s opportunities for entering new markets.     

Tree age influences nutritional,pectin, and anatomical changes in developing ‘Kinnow’ mandarin (Citrus nobilis Lour × Citrus deliciosa Tenora) fruit

An understanding of variation in fruit quality in relation to tree age can help understand the issue of poor fruit quality in young orchards; however, limited information is available on the citrus fruit. In this study, the endogenous nutritional status in rind, rag, and leaves; pectin status in rind and rag; and anatomical fruit growth parameter in rind tissues were studied in ‘Kinnow’ mandarins fruit during their development on trees from three age (6-, 18-, and 35-year-old) groups. In older (35-year-old) trees, rind, rag, and leaf nitrogen (N), phosphorus (P), and potassium (K) concentrations were superior. In fruit from all tree age groups, total pectin and protopectin reduced; however water-soluble pectin (WSP) improved. In rind tissues harvested from young (6-year-old) trees, cell density was more while cell size was less. In all tree age groups, cell density in rind tissues correlated negatively with rind WSP.     

Morphological biodiversity and local use of the Himalayan pear (<Emphasis Type="Italic">Pyrus pashia</Emphasis>) in Central Bhutan

A survey was undertaken in two valleys of the Bumthang area in Central Bhutan to study the morphological diversity, farmers’ knowledge, management practices, yield and income generation potential of the neglected Himalayan Pear (Pyrus pashia Hamilton ex D. Don). For this study a total of 148 Pyrus pashia accessions were collected and 103 households interviewed. Three quantitative and 14 qualitative traits such as colour, size, shape and texture of fruits and tree habit were used for morphological characterization of the accessions. The results demonstrated a high morphological diversity within the P. pashia trees. Fruit shape varied widely from globose to pyriform, fruit skin colour from light yellow and glossy to brown and dull. The observed diversity is likely due to the fact that under farmers’ conditions trees are propagated through seedlings rather than by grafting. None of the characters were specific and correlated with region or altitude. Distinct local cultivars with specific characters were absent making it impossible to group the pears morphologically. Interviews revealed low numbers of trees per farmers, lacking marketability of the fruits and farmers’ subsequent preference for improved varieties as major obstacles for the in situ preservation of biodiversity in P. pashia in this area.     

Reproductive biology and characterization of <Emphasis Type="Italic">Allanblackia parviflora</Emphasis> A. Chev. in Ghana

Allanblackia parviflora A. Chev. (Clusiaceae Lindley) is an indigenous tree species which is found in the rain forest regions of Ghana. It is a potential candidate fruit tree species being introduced in agroforestry systems. Information on genetic diversity, reproductive biology and fruit yield are not known. Field expeditions to seven populations of Allanblackia parviflora in Ghana were undertaken in 2003–2006 during which fruits were collected from 109 trees for characterization. The species was found to be dioecious. The colour of flowers varied from whitish to reddish. No ecological differences in number of fruits per tree, fruit shape and seed health were observed. However, large variations in fruit size and shape were observed among individual trees sampled. A high heritability (h ² = 0.822) in fruit size and genetic gain (G = 20.1%) in fruit size for selection of trees with above average fruit size were observed. A positive significant correlation was observed between fruit size and seed weight (R = 0.54, P < 0.05; Fig. 6). The results suggest that selection and cloning of trees with large fruits could lead to higher yield of seeds for oil production.     

Compound inflorescence cardamom (Elettaria cardamomum (L.) Maton) in India

Cardamom (Elettaria cardamomum (L.) Maton) is an important spice crop of India and valued for its dried capsules. Regular surveys are being undertaken by Indian Institute of Spices Research (IISR), Appangala, Karnataka, India for collecting the variable germplasm of cardamom. At present 436 cardamom accessions collected from cardamom growing tracts of South India are conserved in field gene bank. Among the collections, 23 accessions with varied branching pattern were assembled during explorations and characterized for 16 plant, panicle and capsule characters during 2003–2005. The coefficient of variation was greatest for yield per plant (77.94%) followed by branches per panicle (49.40%) and minimum for dry recovery per cent (7.73%). Two accessions, IC 547214 and IC 349544 were identified as promising with desirable yield contributing characters. The accessions, IC 349553 with extensive branching at the proximal end of the panicle and IC 547180 with dark green capsules even after drying were found to be unique. All the characterized accessions are maintained under ex situ cardamom gene bank of IISR.     

Morphological and molecular diversity of an underutilized fruit crop - <Emphasis Type="Italic">Cordia myxa</Emphasis> L. germplasm from the arid region of Rajasthan,India

Twenty two germplasm accessions of Cordia myxa were collected from Rajasthan and established at the field gene bank for conservation and evaluation. Morphological characterization of 10 year-old trees for 17 traits indicated wide variations among the accessions tested. Higher number of flowers per cyme was found in accession ACHM11 and higher pulp:stone ratio in AHCM25. Overall, AHCM22 was found to be a superior germplasm line for most of the horticulturally useful traits among the accessions tested as it had higher percent of fruit set, pulp:stone ratio and fruit weight. High significant positive correlation was obtained between leaf, fruit characters and pulp:stone ratio. However, these characters were found to be negatively correlated with number of flowers per cyme. Out of 50 random decamer primers used for random amplification (RAPD), 25 were polymorphic. Average polymorphism resolved by these markers among these accessions was 69.8% with an average polymorphic information content of 0.43. Genetic diversity revealed by Jaccard’s co-efficient was between 0.44 and 0.94, and three major clusters were identified among these accessions by phylogenetic analysis using NTSYSpc-2.02e software. RAPD markers associated with leaf size and pulp:stone ratio were also identified. This study shows the existence of high genetic diversity among these accessions.     

Variability in Mucuna pruriens L. germplasm for L-Dopa, an anti parkinsonian agent

Mucuna pruriens L. (Fabaceae) is a popular Indian medicinal plant, which has long been used in traditional Ayurvedic system of Indian medicine for diseases including parkinsonism. The seeds of M. pruriens contain 3,4-dihydroxyphenyl-L-alanine (L-dopa) as a major constituent, an intermediate precursor of the neurotransmitter dopamine. L-dopa extracted from seeds of Mucuna is more effective as drug than the synthetic counterpart. The plant is widely distributed throughout the India and in other parts of the tropics including Central and South America. The present study was aimed to screen thirty-eight accessions of Mucuna germplasm collected from different geographical regions of India for L-dopa content, an effective drug used for Parkinson’s disease. The L-dopa content in seeds of M. pruriens was quantitatively analyzed by high performance thin-layer chromatography (HPTLC) at 280?nm wavelength. Results indicated wide variability among thirty-eight accessions of M. pruriens for seed characters like seed colour, size and shape. The L-dopa content in seeds of M. pruriens ranged between 3.29 and 5.44?% (on dry weight basis) among different germplasm collections augmented from various parts of India. Promising accessions identified having high L-dopa content were IC551549 (5.44?%), IC551514 (5.41?%), IC331536 (5.40?%), IC552857 (5.36?%), IC385844 (5.32?%) and IC439197 (5.32?%). Due to the economic importance of the drug, these promising M. pruriens accessions can be exploited for large scale cultivation for their pharmacological important constituents mainly L-dopa content.     

<Emphasis Type="Italic">Morinda citrifolia</Emphasis> L. (Rubiaceae): a multi-purpose tree for coastal ecosystems and its variability in Konkan region of India

Noni (Morinda citrifolia L.) or Indian mulberry is an evergreen tree species of coastal tropics and has several uses as herbal medicine, cosmetic, vegetable, animal feed, dye and timber. It is a storehouse of useful phytochemicals including caprylic acid, damnacanthal, lineolic acid, morindine, octonic acid and xeronine. The species is hardy, tolerant to shade and salinity hence suitable for marginal lands of tropical coastal ecosystem. Nineteen accessions of noni were collected across the Konkan coast of Goa and Maharashtra states of western India. They are conserved in Indian National Gene Bank assigned with national identity as indigenous collection (IC) numbers viz., IC 0595272 to IC 0595277; IC 0598228 to IC 0598231; IC 0598515 to IC 0598516; IC 0598232 to IC 0598235; IC 0612951 to IC 0612953. The field grown germplasm accessions were characterised using quantitative traits of stem, leaf, flower, fruit and seeds. High degree of variation was recorded for fruit weight and number of seeds per fruit. The results are discussed in the light of evolution, dispersal and utilization. This report aims to summarize the various reports on uses, composition of phytochemcials, and diversity of noni to harness its utilization by increasing human inhabitants in tropical coastal lands.     

Diversity among melon (Cucumis melo L.) landraces from the Indo-Gangetic plains of India and their genetic relationship with USA melon cultivars

We report here the first broad genetic characterization of farmer-developed landraces of melon (Cucumis melo L.) from the Indo-Gangetic plains of India, an area overlooked in previous melon genetic diversity analyses of Indian melon germplasm. Eighty-eight landraces from three melon Groups in two subspecies (C. melo subsp. agrestis Momordica Group, and C. melo subsp. melo Cantalupensis Group and Reticulatus Group) were collected from the four agro-ecological regions (six sub-regions) of two northern states of the Indo-Gangetic plains of India, Uttar Pradesh and Uttarakhand. Significant differences were found among the landraces and eight USA Reticulatus Group reference cultivars for 18 plant and fruit traits: no. of primary branches per plant, days to marketable maturity, sex expression, fruit shape, flesh colour, netting, no. of fruit per plant, fruit weight, shelf life, total soluble solids (°Bx), ascorbic acid (mg/100 g), titratable acidity (%), fruit length and diameter, seed cavity length and diameter, flesh thickness, and resistance to Cucumber mosaic virus. The three melon groups differed significantly for 10 of the plant and fruit traits. Cantalupensis Group and Reticulatus Group accessions were andromonoecious, and the Momordica Group was monoecious. Neighbour-joining (NJ) tree and factorial correspondence analysis (FCA) of simple sequence repeat loci also revealed a high level of genetic variability in this germplasm. The 96 melon genotypes clustered into five groups in the NJ tree analysis: the 16 Indian Reticulatus Group accessions and eight USA reference cultivars formed a distinct group; and the 60 Cantalupensis Group accessions clustered in four other groups with the 12 Momordica Group accessions in a distinct subgroup of one of the Cantalupensis groups. The FCA plot largely confirmed the NJ tree with three distinct groups, one for each melon group. The close affinity of the Indian and USA Reticulatus melons was not unexpected, but it is not clear whether it was inherent in the group and maintained as Reticulatus melons moved from India through Central Asia and Europe to North America, or the result of recent intercrossing of Indian landraces with the USA-derived cultivars and selection for a broad range of Reticulatus type melons.     

Differences in quality parameters and nutrient composition in Fukumoto oranges with and without creasing symptoms

Creasing is a physiological disorder in oranges fruits that affects rind quality. This disorder has been associated with insufficient or imbalances of nutrients in the fruit. A study was carried out in an orange orchard cv. Fukumoto. At harvest, fruit samples were selected with and without creasing, according to visual symptoms. External and internal fruit quality was evaluated in whole fruit, then dry matter and mineral content and concentration (nitrogen N, potassium K, calcium Ca and magnesium Mg) was analyzed in flavedo, albedo and pulp. Oranges with creasing had larger weight and fruit size, higher water content in the pulp and whole fruit, lower coarse rind grading, thinner peel, higher nitrogen content in the pulp and the whole fruit, higher potassium content in all fruit components and lower calcium content and concentration. Magnesium content, only in flavedo showed significant differences, being higher in fruit without creasing. Fruit size and Ca were key factors related to the expression of creasing disorder, as well as nutrient balance.     

Morphological variability and resource allocation in the annual teosinte (<Emphasis Type="Italic">Zea mays</Emphasis> ssp.<Emphasis Type="Italic"> mexicana</Emphasis> (Schrader) Iltis) of the Toluca Valley,Mexico

This study explores the morphological variability and resource allocation of weedy, annual teosinte (Zea mays ssp. mexicana (Schrader) Iltis, Chalco race) from the Valley of Toluca, central Mexico, and compares it with data reported for maize. One hundred mature and fertile teosinte plants were selected for variation in size and measured. The extremes found for fertile plants were 0.12 and 3.5 m in height and 0.09 vs. 694 g in weight. Fruit were polymorphic, with significant correlation between color and weight. Small plants invested slightly more biomass in the leaves and tall plants invested more in the stem. The minimum mass necessary to produce fertile plants was much lower in teosinte than in maize. Very small teosinte plants formed only female inflorescences, whereas maize produces a tassel. The harvest index was similar to landrace maize, and did not decrease in taller plants, as teosinte can increase the reproductive sink. Toluca teosinte retains several important characteristics of weeds: size variability, adaptation of fruit quantity to environmental conditions, and fruit polymorphism.     

Genetic diversity analysis in Boerhavia diffusa L. of different geographic locations in India using RAPD markers

Boerhavia diffusa is extensively used in herbal medicines as well as in the Ayurvedic system, because it contains a set of clinically important compounds. In the present study, the genetic variability in Boerhavia diffusa between accessions of different geographical origin within the Indian territory is assessed through random amplified polymorphic DNA (RAPD) markers. Twenty-eight accessions of Boerhavia were screened with eighteen primers of which nine were found to be the most informative. The degree of polymorphism was found to be high in accessions collected from different places of Uttar Pradesh (Set II) in comparison to other states of India (Set I). A relatively lower level of polymorphism was recorded in accessions collected from diverse locations around Lucknow (Set III). Accessions from neighbouring geographical regions exhibited more similarity than those from distant regions (as revealed by the set I analysis). Certain diagnostic markers may be correlated with morphological character(s) such as plant type. BDL appeared most distinct and divergent from the rest of the accessions and the BDJ plant in set II also showed least similarity estimate. Fragments of 5.62 Kb and 4.47 Kb with primer GN59 was found to be unique for BDP and BD2 having ovate leaf character, whereas ovoid leaf genotype exhibited 0.79 Kb (GN34 primer) fragment. Similarly a unique band type (0.35 Kb) with primer GN83 was present in BDL and BD1 that share light pink flower. Jaccard's, and Nei and Li similarity coefficient values amongst the accessions were in the range of 0.22 to 0.89 and 0.33 to 0.93, respectively. Association of RAPD markers with the leaf characteristics, flower colour as well as with geographical locations has been made. This shows that RAPD markers are also useful for the study of genetic structure of Boerhavia populations.     

A wide range of genetic diversity in pear (<Emphasis Type="Italic">Pyrus ussuriensis</Emphasis> var. <Emphasis Type="Italic">aromatica</Emphasis>) genetic resources from Iwate,Japan revealed by SSR and chloroplast DNA markers

Iwateyamanashi (Pyrus ussuriensis var. aromatica) is one of the Pyrus species which grows wild in Japan. The number of Iwateyamanashi trees has been decreasing, so conservation and evaluation is urgently needed. Over 500 accessions of Pyrus species collected from Iwate in northern Tohoku region are maintained at Kobe University as an Iwateyamanashi germplasm collection. In order to investigate the genetic diversity, five SSR (simple sequence repeat) markers, developed from Japanese and European pear were examined for 86 Pyrus individuals including 58 accessions from Iwate. These SSR loci could discriminate between all the Iwate accessions except for 10 that bear seedless fruit, as well as determine the genetic diversity in Iwateyamanashi germplasms. High levels of variation were detected in 41 alleles and the mean observed heterozygosity across 5 loci was 0.50 for the Iwate accessions. Seedless accessions sharing identical SSR genotype with the local pear variety “Iwatetanenashi” were supposed to have been propagated vegetatively via grafting. In an UPGMA phenogram, Japanese pear varieties (P. pyrifolia) were clustered into two groups with some Iwate accessions including seedless ones. Another 38 Iwate accessions were not clustered clearly, and there was no clear relationship between these accessions and geographical distribution or morphological characters. Allele frequency revealed that the Iwate accessions were genetically more divergent than the Japanese pear varieties. Most Japanese pears possessed a 219 bp deletion at a spacer region between the accD and psaI genes in the chloroplast DNA (cpDNA), but other Pyrus species and two Iwateyamanashi trees did not. In the Iwate accessions, 79.3% had a deletion type cpDNA and others had a standard type cpDNA without deletion. These results are indicative of the wide range of genetic diversity in the Iwate accessions which include Japanese pear varieties. A combination of SSR and cpDNA analyses revealed high heterogeneity in Iwateyamanashi and coexistence of Iwateyamanashi and hybrid progeny with P. pyrifolia. These could be reasons for the wide range of continuous morphological variation described previously.     

Phenotypic diversity within native Iranian almond (<Emphasis Type="Italic">Prunus</Emphasis> spp.) species and their breeding potential

A total of 137 accessions from 18 wild almond species were collected from Iran and leaf and fruit traits were characterized. Also evaluated were flowering and ripening date, self-incompatibility and kernel bitterness. An extensive phenotypic diversity was found both among and within species. Differences in average leaf dimensions among and within species were associated with average rainfall but not altitude of collection site. Adjacent accessions located in drier areas had smaller leaf dimensions than those located in semi-humid or humid regions. No relation was found between average fruit dimensions and collection site conditions. Principal component analysis revealed that the nut weight and width, and the kernel weight had highest loading in the first component accounting for 45.8% of total variation. In contrast, leaf traits in the second component accounted for 22.3% of total variation. No significant correlations were detected between leaf dimensions and fruit traits in all species evaluated. Results document a rich source of new germplasm for almond improvement programs. Small fruit size, pollen-pistil self-incompatibility, and bitter kernel flavour are the most common obstacles to the utilization of this wild germplasm in breeding.     

Agro-morphological diversity of Spanish traditional melons (<Emphasis Type="Italic">Cucumis melo</Emphasis> L.) of the Madrid provenance

The variability of 14 landraces belonging to a Madrilean village historically specialized on melon cultivation, Villaconejos, was evaluated based on 58 quantitative and qualitative morphological traits. These landraces were compared to a reference array composed of 14 accessions which represented the main varieties cultivated in Spanish fields. Individual data related to plant, fruit, seed and phenology have been analysed using a multivariate analysis. This analysis showed intra- and inter-varietal diversity, as emphasized the most discriminant morphological traits in order to define similarities. Villaconejos accessions were morphologically distinct from the reference accessions (RA). Only the landraces belonging to Piel de Sapo market class showed common morphological affinities with the RA, grouping with them. The majority of Villaconejos traditional varieties clustered in five different groups, with no RA within, showing distinctive morphological singularities not described previously. Two of these groups showed some particularities in fruit traits, which are appreciated as quality marks for Spanish consumers. These results indicate that these accessions must be conserved as valuable genetic resources to enrich the Inodorus genetic bases for future breeding proposes worldwide. Furthermore, it should be considered the opportunity of promoting their cultivation under the shelter of a Protected Geographical Indication as a high quality melon. Finally, the discovering of such a high variability presented in a very small area, gives a clue for focusing, with an elevated probability of success, future surveys in similar ancestral European villages which in the past times, also supplied more populated cities with their farming products.     

Characterization of a landrace collection for Tomàtiga de Ramellet (Solanum lycopersicum L.) from the Balearic Islands

Tomàtiga de Ramellet (Ramellet) is a traditional long-storage tomato from the Mediterranean Islands of Mallorca, Menorca, Eivissa (Ibiza), and Formentera (Balearic Islands). To investigate the observed phenotypic and shelf life variability within local varieties, we studied 142 accessions characterized as Ramellet varieties, and 29 accessions representing other local varieties. The 171 accessions constitute a new germplasm collection at the University of Balearic Islands (UIB). The evaluation of the collection demonstrated wide variability in most agronomic, morphological and quality traits of Ramellet accessions and clearly differentiates them from other local varieties with respect to fruit shape, size, fruit composition, and fruit storage. All accessions of Ramellet evaluated have the ripening mutation alcobaça (alc), suggesting an important role for this variant in controlling shelf life. Results of evaluation also showed wide variability between the Ramellet accessions for storage time, suggesting that other traits like fruit size and composition may play a role in fruit conservation. Observed variation within the Ramellet accessions is consistent with a genetically variable population or landrace group. The establishment of this collection will help prevent genetic erosion of this local landrace group and constitute a resource for future investigation and breeding.     

Phenotypic and genomic characterization of vine cactus collection (Cactaceae)

Hylocereus (Berger) Britton et Rose, Selenicereus (Berger) Britton et Rose and Epiphyllum Haw. species have commercial potential as exotic fruit crops in semi-arid and arid lands. The high genetic variability among these species offers an opportunity for commercial cultivation. Toward this end we investigated genomic and morphological characteristics including: nuclear DNA content (2C-values), stomatal length and density, potential yield and reproductive parameters in 64 Hylocereus, Selenicereus and Epiphyllum accessions. Nuclear DNA content ranged from 3.21 pg for S. grandiflorus (L.) Britton et Rose spp. grandiflorus to 8.77 pg for H. megalanthus (Vaup.) Bauer. All species were diploid except the tetraploids H. megalanthus and S. vagans (Bgek.) Britton et Rose. Stomatal length and density, fruit weight, potential yields, number of viable seeds per fruit and fruit maturation times were highly variable among accessions. No significant correlations were found between stomatal length, density, and nuclear DNA content, nor between fruit weight and seed number. The high genetic variability found between the accessions here provides further support for the excellent prospects of conserving and domesticating these exotic species.     

Changes in seed Morphology, Dormancy and Germination from wild to Cultivated Hyacinth bean Germplasm (Lablab purpureus: Papilionoideae)

Morphological and physiological seed characteristics of the hyacinth bean (Lablab purpureus) were investigated in a set of 18 different germplasm accessions, from wild over semi-domesticated forms to landraces and current cultivars. The objective of this study was to improve the understanding of the domestication of this tropical legume crop. Wild accessions were easily distinguished by morphological characters, such as small, greyish-brown, mottled seeds. Cultivated or semi-domesticated forms showed a much wider variation in size, colour and shape. Seed mass of cultivated accessions reached up to almost 10 times that of wild accessions. Most wild accessions showed a greater spread of germination and larger proportions of hard seeds, up to about 70%, than most cultivated and so-called semi-domesticated accessions. Cluster analysis applied separately to morphological and germination data and a subsequent discriminant analysis did not help integrating the morphological variability observed. The presumed occurrence of semi-domesticated accessions within the germplasm tested as well as the crop's origin from Africa alone or both Africa and Asia are debated on the basis of seed germination and hard seed coat dormancy.