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

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

Water Relations of Seed Development and Germination in Muskmelon (Cucumis melo L.) : IV. Characteristics of the Perisperm during Seed Development

We previously reported that an apparent water potential disequilibrium is maintained late in muskmelon (Cucumis melo L.) seed development between the embryo and the surrounding fruit tissue (mesocarp). To further investigate the basis of this phenomenon, the permeability characteristics of the tissues surrounding muskmelon embryos (the mucilaginous endocarp, the testa, a 2- to 4-cell-layered perisperm and a single cell layer of endosperm) were examined from 20 to 65 days after anthesis (DAA). Water passes readily through the perisperm envelope (endosperm + perisperm), testa, and endocarp at all stages of development. Electrolyte leakage (conductivity of imbibition solutions) of individual intact seeds, decoated seeds (testa removed), and embryos (testa and perisperm envelope removed) was measured during imbibition of freshly harvested seeds. The testa accounted for up to 80% of the total electrolyte leakage. Leakage from decoated seeds fell by 8- to 10-fold between 25 and 45 DAA. Presence of the perisperm envelope prior to 40 DAA had little effect on leakage, while in more mature seeds, it reduced leakage by 2- to 3-fold. In mature seeds, freezing, soaking in methanol, autoclaving, accelerated aging, and other treatments which killed the embryos had little effect on leakage of intact or decoated seeds, but caused osmotic swelling of the perisperm envelope due to the leakage of solutes from the embryo into the space between the embryo and perisperm. The semipermeability of the perisperm envelope of mature seeds did not depend upon cellular viability or lipid membrane integrity. After maximum seed dry weight is attained (35-40 DAA), the perisperm envelope prevents the diffusion of solutes, but not of water, between the embryo and the surrounding testa, endocarp, and mesocarp tissue.     

Water entry in Cercis siliquastrum (Leguminosae) seeds

By use of scanning electron and light microscopy a study on the histological organization controlling water entry in the Cercis siliquastrum L. seeds was carried out. The intact mature dry seed of Cercis is impermeable. As we have previously demonstrated, this property is prevalently due to a non–cellular layer situated at the inner border of the testa enclosing the endosperm. When this layer is damaged, water enters endosperm and embryo, increasing their volume, and stretching the testa. The testa copes with the increasing volume of the seed without rupturing and decreases in thickness. The shape of the hypodermal cells changes from almost circular in section to periclinally elongated, as an effect of the imbibition.     

Relation Between the Anatomy of the Testa, Water Permeability and the Presence of Phenolics in the Genus Pisum

The seed coats of Pisum elatius, P. fulvum and P. humile areimpermeable to water while those of P. sativum and P. humilex P. sativum are permeable. The anatomical structure of theseed coats and the location of phenolics and quinones in thecells is described. The barriers to permeation of water in theimpermeable seeds are the continuous, very hard, pectinaceouslayer of the caps of the palisade cells and the presence ofquinones in either the palisade or osteosclereid cells, in acontinuous layer of these cells. In water permeable seeds thecaps are looser and quinones discontinuous or absent in palisadeor osteosclereid cells. Pisum, testa, water permeability, quinones, phenolics, palisade cells, osteosclereids, pectinaceous caps     

Role of seed coat in imbibing soybean seeds observed by micro-magnetic resonance imaging

Background and Aims

Imbibition of Japanese soybean (Glycine max) cultivars was studied using micro-magnetic resonance imaging (MRI) in order to elucidate the mechanism of soaking injury and the protective role of the seed coat.

Methods

Time-lapse images during water uptake were acquired by the single-point imaging (SPI) method at 15-min intervals, for 20 h in the dry seed with seed coat, and for 2 h in seeds with the seed coat removed. The technique visualized water migration within the testa and demonstrated the distortion associated with cotyledon swelling during the very early stages of water uptake.

Key Results

Water soon appeared in the testa and went around the dorsal surface of the seed from near the raphe, then migrated to the hilum region. An obvious protrusion was noted when water reached the hypocotyl and the radicle, followed by swelling of the cotyledons. A convex area was observed around the raphe with the enlargement of the seed. Water was always incorporated into the cotyledons from the abaxial surfaces, leading to swelling and generating a large air space between the adaxial surfaces. Water uptake greatly slowed, and the internal structures, veins and oil-accumulating tissues in the cotyledons developed after the seed stopped expanding. When the testa was removed from the dry seeds before imbibition, the cotyledons were severely damaged within 1·5 h of water uptake.

Conclusions

The activation of the water channel seemed unnecessary for water entry into soybean seeds, and the testa rapidly swelled with steeping in water. However, the testa did not regulate the water incorporation in itself, but rather the rate at which water encountered the hypocotyl, the radicle, and the cotyledons through the inner layer of the seed coat, and thus prevented the destruction of the seed tissues at the beginning of imbibition.Key words: Dry seeds, Glycine max, MRI, seed coat, soaking injury, soybean, testa, role of inner layer of seed coat, water uptake     

The Damaging Effect of Water on Dry Pea Embryos During Imbibition

When pea seeds were imbibed in water without their seed coats,vital staining revealed that cells on the abaxial surface ofthe cotyledons were dead. No damage occurred on the surfaceof cotyledons when the seeds were imbibed intact, or beneaththe testa when only half of the testa was removed. Cell deathoccurred as a result of rapid water uptake within the first2 min of imbibition, since reducing the rate of imbibition insolutions of Carbowax 4000 lessened the damage. Cell death wasrestricted to the outer layers of the cotyledons; inner tissuesremained alive. These observations supported the hypothesisthat rapid early leakage during imbibition of dry embryos resultedfrom the death of cells caused by the physical disruption ofmembranes. Imbibition damage resulted in reduced respirationand germination, a decline in the rate of food reserve transferfrom the cotyledons to the growing axis, and a lower growthrate in the seedlings produced. Greater sensitivity of embryosto imbibition damage at low temperature, and similarities betweenfeatures of imbibition damage and chilling injury led to thesuggestion that so-called chilling injury is the result of imbibitiondamage rather than the effects of low temperature.     

Leaching of Ions, Organic Molecules, and Enzymes from Seeds of Peanut (Arachis hypogea L.) Imbibing without Testas or with Intact Testas

Reducing sugars, phosphates, potassium ions, total electricallyconducting material, proteins, and phenolics leached from twovarieties of peanut (Arachis hypogea L. ) seeds during 24 himbibition in distilled water. The same substances leached fromthe seeds when testas were removed before imbibition. The quantitiesof substances leached from seeds without testas after 24 h,except protein and phenolics, were much greater than from seedswith intact testas. Time courses of leaching of sugars, phosphates,potassium ions, and of total electrically conducting materialshowed fastest leaching rates in the first hour of imbibition.In the first 4 h of imbibition more sugars leached from intactseeds than from seeds without testas. Peroxidase was presentin leachates from intact seeds but was undetectable in leachatesfrom seeds without testas. Hydrogen peroxide-reducing activitywas detected in 24 h leachates from intact seeds and from seedswithout testas. The activity was partly inhibited by 1 mM ascorbate.It was concluded that catalase was present in the leachates. The peanut testas were thin. Their presence around the embryodid not reduce the rate of imbibition as compared with thatof seeds without testas. Thus the greater leaching of some substancesin the absence of the testa could not be ascribed to the absenceof a physical barrier to water uptake.     

Cauliflower (Brassica oleracea L.) Seed Vigour: Imbibition Effects

By means of a machine vision facility, the process of water-imbibitionin a small seeded brassica species was recorded as the visibleincrease in seed volume. Dry cauliflower seed (Brassica oleracea)showed an immediate rapid phase of imbibition upon the additionof water. This initial phase was associated with the reductionin seedling root growth resulting from the imbibition of coldwater, and the rate of uptake of water at 20?C was negativelycorrelated with subsequent seedling growth. Damage to the testaof dry seed resulted in an increased rate of imbibition anda corresponding decrease in seedling root growth measured onslant boards. This showed that the intact testa of cauliflowerseed is capable of acting as a barrier to water influx and thata high rate of water uptake is damaging to the embryo. Testadamage reduced percentage soil emergence of seeds, as did raisingthe soil moisture content during the imbibition period by wateringimmediately after sowing. Both these treatments were believedto increase imbibition rate. Conditions which encouraged a lowrate of water uptake also improved the rate and uniformity ofemergence. Correlation of mean imbibition rates (measured ina laboratory test) with soil-emergence indicated that the sensitivityto imbibition damage varied between seed lots and interactedwith the absolute rate of water influx to determine the finalpercentage emergence. This factor prevented reliable predictionof seedling performance from the ranked-order of measured imbibitionrates. The significance of these findings to the seed productionand modular transplant raising industries is discussed. Key words: Machine vision, imbibition rate, testa, soil moisture, vigour     

DEVELOPMENT,STRUCTURE AND FUNCTION OF SUBPALISADE CELLS IN WATER IMPERMEABLE SIDA SPINOSA SEEDS

Newly matured prickly sida (Sida spinosa) seeds were hard, but afterripening, heat or pressure permitted water entry solely via a predefined region of the chalazal area. In this region, the raising of a “blister” formed by separation of the palisade of the seed coat from underlying tissues preceeded measurable water uptake by prickly sida seeds. A single subpalisade layer, unique to the region beneath the blister, was involved in the sequence of events in seed water uptake. The lateral walls of subpalisade cells invariably broke near the palisade border and cell contents were extruded onto the surface. This report describes the cytological development of the subpalisade layer from 1-21 days post anthesis. Cells beneath the potential “blister” near the chalazal slit developed into columnar subpalisades and cells beneath the subpalisades or beyond the margins of the potential blister, developed into oval, thick-walled chalazal cap cells. By 6-10 days, distinct features of the subpalisades included: 1) thin portions in lateral walls due to lack of secondary wall depositions at the palisade border; 2) progressive accumulation of fibrous material in numerous vacuoles; and 3) progressive coalescence of osmiophilic bodies and degeneration of cytoplasmic contents. At 21 days, the seeds were dehydrated, mature and hard, but the thin, lateral subpalisade walls were still intact and had not broken. The thin-walled portions were predetermined weak sites that break, permit palisade separation, expose the area under the blister to available moisture and result in subsequent imbibition of water by the seed. The hydrophilic, fibrous material extruded from the ruptured subpalisade cells may attract water to the newly exposed surface and facilitate penetration of water into the nutritive and embryonic seed tissues.     

Seed after-ripening and over-expression of class I beta-1,3-glucanase confer maternal effects on tobacco testa rupture and dormancy release

'Coat-imposed' seed dormancy of many non-endospermic and endospermic species is released during after-ripening. After-ripening-mediated promotion of tobacco ( Nicotiana tabacum L.) seed germination is mainly due to a promotion of testa rupture and a similar promotion of subsequent endosperm rupture. Treatment of after-ripened or freshly harvested mature seeds with abscisic acid (ABA) delays endosperm rupture and inhibits the induction of class I beta-1,3-glucanase (betaGlu I) in the micropylar endosperm, but does not affect the kinetics of testa rupture. After-ripening-mediated release of photodormancy is correlated with a decreased gibberellin (GA) requirement for testa rupture during dark-imbibition. Reciprocal crosses between wild-type tobacco and sense-betaGlu I transformant lines showed that betaGlu I over-expression in the seed covering layers can replace the promoting effect of after-ripening on testa rupture in light, but only if the mother plant is a sense-betaGlu I line. This maternal effect supports the model of two sites for betaGlu I action: (i) betaGlu I contribution to the after-ripening-mediated release of dormancy in the dry seed state, which is manifested in the promotion and ABA-insensitivity of testa rupture during imbibition. (ii) ABA-sensitive expression of betaGlu I in the micropylar endosperm, which contributes to endosperm rupture. The importance of GA-signaling and testa characteristics appear to be a common feature during the after-ripening-mediated release of coat-imposed dormancy in endospermic and non-endospermic seeds.     

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

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

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

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

濒危植物中甸刺玫种子休眠及其生态学意义

对中甸刺玫Rosa praelucens种子结构及其透水性,赤霉素处理对胚的影响,果壳、种皮、胚乳的粗提物活性进行研究。结果表明：(1) 中甸刺玫外种皮是由多层排列紧密的厚壁细胞组成,内种皮为坚硬致密的栅栏组织;(2) 种皮对种子的吸胀阻碍较大, 未处理的种子吸水率较低,吸水13 d后增加量为18.82%;(3) 果实结籽率为0.69%,多数果实中没有饱满的种子;(4) 赤霉素100 mg·kg-1预处理种子可加快胚的萌发速率;(5) 胚乳、种皮、果壳中存在内源抑制物。中甸刺玫种子的休眠是由其形态和生理特点引起的综合休眠。     

Magnoliid reproductive organs from the Cenomanian-Turonian of north-western Kazakhstan:Magnoliaceae andIlliciaceae

Cenomanian-Turonian sediments from the Sarbay locality in north-western Kazakhstan have yielded a rich assemblage of plant fossils including well preserved flowers, fruits, and seeds of angiosperms. This work describes fossil seeds assigned to theMagnoliaceae and theIlliciaceae. Three new species of the extinct magnoliaceous genusLiriodendroidea, L. asiatica, L. costata, andL. tenuitesta, are established and new information on the previously described species,L. alata, is provided. TheLiriodendroidea seeds are closely similar to seeds of extantLiriodendron, but are distinguished in being much smaller and winged. A new genus and species,Illiciospermum pusillum, is established based on seeds with close similarity to those of the extant genusIllicium. The seeds are small, anatropous and exotestal with outer epidermis of testa forming a palisade layer. The facets of the palisade cells have deeply undulate anticlinal walls. The micropyle area is seen on the outer integument as a transverse slit placed on a raised strophiole-like structure close to the hilum. TheIlliciospermum seeds represent the first unequivocal record of theIlliciaceae in the Cretaceous. Another seed of possible illiciaceous affinity is described as aff.Illiciospermum sp.     

The Role of the Lens in Seed Imbibition and Seedling Vigour of Sesbania punicea (Cav.) Benth. (Leguminosae: Papilionoideae)

Seeds of Sesbania punicea (Cav.) Benth. are largely impermeable,and the natural, low, percentage which germinate are permeableat the lens. Scarifying seed in acid stimulates germinationand causes various types of damage to the testa, including thelens. Placing seeds in boiling water stimulates germinationby rendering the seeds permeable at the lens. Seeds were mechanicallyscarified at precise sites on the testa to assess influenceof the site and degree of damage on germination. Results indicatethat both the site of initial water entry and the rate of waterentry are important in determining seed and seedling vigour.Damaging the lens is more effective in stimulating germinationand reducing seedling abnormalities than complete excision ofthe lens. This effect can be reversed if seeds from which thelens was excised are allowed to imbibe gradually in inert osmotica.The leguminous lens thus acts as the site of initial water entryand regulates the rate of water uptake, thereby increasing seedand seedling vigour. Sesbania punicea (Cav.) Benth., hardseededness, lens, seed germination, imbibition damage, seedling vigour     

Water impermeable seed dormancy

Viable seeds that do not imbibe water and thus fail to germinate in an apparently favorable environment are commonly termed impermeable or hard seed. This physical, exogenous dormancy is especially common in species of the Fabaceae. The ecological significance of hard seed includes the ability to rapidly recolonize burnt areas after fire and to withstand ingestion by animals and birds. Advantages and problems that hard seed cause in agriculture are discussed. Species from different families with impermeable seeds appear to have in common a layer of macrosclerid cells that form a palisade layer in the testa. The term strophiole and its contradictory use in botanical literature are discussed. Genetic factors and environmental conditions both affect the proportion of impermeable seeds produced. Methods of artificially softening impermeable seeds include acid and solvent, soaking, mechanical scarification, pressure, percussion, freezing, heating, and radiation treatments that can result in a change in germination from less than 20% in some untreated species up to 90% or more in treated species. Natural softening involves high temperatures and temperature fluctuations and the degree of desiccation of the seed. The mechanism of water impermeability is related to the testa and is thought to involve waterproofing substances including wax, lignin, tannin, suberin, pectin, and quinone derivatives. The hilum acts as a hygroscopic valve that prevents water uptake but allows water loss to occur at low relative humidities in some species. The strophiole is an area of weakness in the testa of some Papilionoideae while the chalaza region has been determined as an area of weakness inPisum andGossypium. The water impermeable status of some species is reversible at a seed moisture content greater than 10%. The hard seed of a species can be described both in terms of the amount and the degree of impermeability.     

A Seed Shape Mutant of Arabidopsis That Is Affected in Integument Development

A seed shape mutant of Arabidopsis was isolated from an ethyl methanesulfonate-treated population. Genetic analysis revealed that the heart-shaped phenotype was maternally inherited, showing that this is a testa mutant. This indicated the importance of the testa for the determination of the seed shape. This recessive aberrant testa shape (ats) gene was located at position 59.0 on chromosome 5. A comparison was made between ovules and developing and mature seeds of the wild type and of the mutant using light and scanning electron microscopy. We showed that the mutant seed shape is determined during the first few days after fertilization, when the embryo occupies only a very small part of the seed. The integuments of ats ovules consisted of only three rather than five cell layers. In double mutants, the effect of ats was additive to other testa mutations, such as transparent testa, glabra (ttg), glabrous2 (gl2), and apetala2 (ap2). The ats mutation resulted in a reduced dormancy, which was maternally inherited. This effect of a testa mutation on germination was also seen in ttg seeds, in which the outer layer of the testa was disturbed. This indicated the importance of the testa as a factor in determining dormancy in Arabidopsis.     

Growth and curvature in seedlings of Pisum sativum and an ageotropic mutant

In an attempt to explain the influence of gravity on the behaviour of ageotropic plant organs, a pea mutant (Pisum sativum ageotropum) and normal pea (Pisum sativum cv. Sabel) were examined. The mutant has a significantly lower germination rate (large seeds: 25%, small seeds: 10%) than normal pea seeds (55%). Removal of testa increased germination dramatically, the values obtained were 63 and 89%, respectively. Immediately after imbibition the mutant from which the testa had been removed, developed more slowly than normal pea seeds; after 28 h the difference in elongation rate between the two types was reversed. When continuously stimulated geotropically in the horizontal position the elongation in the mutant is larger than in the normal pea roots kept in the same position. During a 24 h period starting 48 h after imbibition the mutant root elongated 45.0 mm while the value for the normal pea root was 11.5 mm. The course of the geotropic curvature in roots of the two types has been followed during a period of 24 h. Normal pea roots develop an asymmetry in the extreme root tip region after 30 min of horizontal stimulation. After prolonged stimulation (exceeding 2 h) the asymmetry has disappeared and the curvature distributed over the entire growth region. When roots of normal pea are stimulated continuously at various angles, the optimum angle of geotropic response is 90° with decreasing responses in the order 135° (i.e. the root tip is pointing obliquely upward) and 45°. The presumed ageotropic behaviour of the mutant has only to a certain extent been confirmed in the present study. When stimulated at 135° a slight positive curvature developed; stimulation at 90° and 45° gave a slight negative curvature.     

Seed morphology and endosperm structure of selected species of Primulaceae, Myrsinaceae, and Theophrastaceae and their systematic importance

Seed size and shape, seed coat surface pattern, seed coat thickness, and endosperm structure were investigated in Androsace septentrionalis, Cortusa matthioli, Hottonia palustris, Primula elatior, Soldanella carpatica (Primulaceae), Anagallis arvensis, A. minima, Cyclamen purpurascens, Glaux maritima, Lysimachia nemorum, L. vulgaris, Trientalis europaea (Myrsinaceae), and Samolus valerandi (Theophrastaceae). Three seed size categories were distinguished on the basis of biometric measurements. Almost all seeds examined were found to be small with an angular shape classified as sectoroid or polyhedral. A new type of seed shape, suboval, was identified for H. palustris. Cyclamen purpurascens seeds differed from seeds of all other species examined because of their large size, subglobose shape, and concave hilar area. Despite the different shape types, the length/width ratio of the seeds examined was constant, while their hilum length/width ratio was a highly variable feature. Three types of seed surface patterns were observed: (1) reticulate, (2) tuberculate with secondary striations, and (3) poroid-alveolate with the presence of a spongy outer layer. For seeds of Anagallis arvensis, A. minima, Cortusa matthioli, Lysimachia nemorum, and Soldanella carpatica, secondary seed sculpture was described. The seed coats of all species examined were two-layered, and great differences in testa thickness were found (9.9?C128.6???m). In general, seeds of the Myrsinaceae species were more often characterized by thick testa. Different proportions in thickness of the inner and outer testa layers were observed. In seeds with reticulate seed patterns, the inner testa layer was twice to several times thicker than the outer layer, while the opposite proportions were observed in seeds with the tuberculate or poroid-alveolate seed sculpture pattern. In seeds of all species examined, oxalate crystals were present on the surface of the inner testa layer. Thus, the presence or absence of oxalate crystals in testa is not a feature distinguishing species with angular or subglobose seeds. Four types of endosperm structure were identified according to the thickness of the endosperm cell walls and the relief of their inner surface: (1) with evenly thickened and smooth cell walls, (2) with evenly thickened cell walls and circular or helical thickenings on their inside surfaces, (3) with very thick, but not evenly thickened cell walls with constrictions (??pitted??), and (4) with very thin papery and undulate cell walls. There is no rule concerning the seed shape, type of the seed sculpture, testa thickness, or endosperm structure that corresponds to the family affiliation of the species examined.     

Seed Structure in Cannaceae: Taxonomic and Ecological Implications

The ovules and seeds of Canna show some striking differencesto those in other zingiberalean families. The pachychalazaldevelopment of the ovule results in a seed of which only a smallpart of the testa is of tegumentary origin. A silicified endotesta,characteristic of the order, is lacking. The mechanical layerof the seed is formed by a continuous exotesta of Malpighiancells. The intact seed coat is impermeable. The seed is ableto absorb water after the raising of a preformed imbibitionlid on the raphe. During imbibition the extotesta loses itshardness, allowing the embryo to emerge. The special structuralfeatures of the seed are discussed in relation to records onthe extreme longevity of Canna seeds. Canna tuerckheimii, C. jaegeriana, C. glauca, pachychalaza, Malpighian cells, imbibition lid, seed longevity