“OBLIGATE” ALGAL EPIPHYTE: SMITHORA NAIADUM GROWS ON A SYNTHETIC SUBSTRATE1

The supposedly obligate epiphyte Smithora naiadum, normally found only on the sea grasses Phyllospadix scouleri and Zostera marina, grew on an artificial substrate under field conditions. This development shows that the alga does not appear to require a chemical factor from its normal host plant for (1) substrate selection and adhesion, (2) growth of basal cushion and young blades, or (3) maturation of thick cushions.     

Negative phototropism in young gametophytes of the fern Schizaea pusilla

Schizaea pusilla is a rare fern that occurs in acidic bogs and is one of the few fern species that maintains a filamentous gametophyte throughout its development. To expand our knowledge of the physiology of this fern, phototropic responses were examined in young gametophytes. In contrast to germ filaments of other fern species, apical protonemata of young gametophytes are negatively phototropic in continuous white, red and blue light at all fluence rates tested. The expression of phototropic curvature is not limited by time since apical protonemata are also negatively phototropic when they are given brief exposures of light and then placed in the dark. In other lower plant groups such as mosses and some algae, the direction of phototropic curvature can change depending on light quality and intensity, but in young gametophytes of Schizaea, negative phototropic curvature was observed in all conditions studied. Blue light is the most effective in promoting the negative phototropic response in Schizaea.     

PHOTOINHIBITION AND RECOVERY AFTER HIGH LIGHT STRESS IN DIFFERENT DEVELOPMENTAL AND LIFE-HISTORY STAGES OF LAMINARIA SACCHARINA (PHAEOPHYTA)1,2

The capacity to cope with high light stress was investigated in different life-history and developmental stages of Laminaria saccharina Lamour. sporophytes and gametophytes. Changes in photosynthetic efficiency and in the level of photoinhibition were measured by in vivo fluorescence changes of photosystem II. Pigment content was studied using high performance liquid chromatography. Additionally, the morphology of the various developmental stages during the life cycle was studied by light microscopy in relation to the photosynthetic parameters. High light stress (2 h, 500 μmol.m-².s^?1) induced photoinhibition of photosynthesis with fast kinetics in older sporophytes and gametophytes. In contrast, the absolute degree of photoinhibition after light stress was higher in younger than in older sporophytes. Photosynthesis recovered faster in older sporophytes and gametophytes compared to young sporophytes. In very young sporophytes, photosynthesis did not recover fully even after 12 h exposure to low light, indicating severe photodamage. Kinetics of recovery in old sporophytes and in gametophytes showed a fast and a slow phase, whereas younger sporophytes recovered only with a slow phase, The fast phase is indicative of a decline of the photoprotective process, whereas the slow phase indicates a recovery from photodamage. The capacity to cope with high light stress in Laminaria sporophytes increased with increasing age of the thalli. The gametophytes are less sensitive to high light stress and may be selected to endure unfavorable white light conditions. Investigation of the xanthophylls showed that the higher resistance to high light is not caused solely by a higher content of xanthophyll cycle pigments. Additionally, changes in the thallus structure during the development of the sporophytes seemed to cause a higher resistance to high light. The observed changes in the ability to cope with high light in the different life-history and developmental stages of Laminaria saccharina may influence the distribution of the species on the shore.     

GAMETOPHYTES AND YOUNG SPOROPHYTES OF OPHIOGLOSSUM CROTALOPHOROIDES WALT.

The lens-shaped, dorsiventral gametophytes of Ophioglossum crotalophoroides are superficially different from the cylindrical gametophytes of other species of Ophioglossum. However, they have the same features as other Ophioglossum gametophytes (fundamentally axial organization, radially symmetrical apical meristem, radial distribution of gametangia) except that the cylindrical axis is reduced in length. Young sporophyte development is unique in the genus: all primary organs clearly arise from the embryo and develop simultaneously. The length of the life cycle of O. crotalophoroicles is considerably shorter than that of some other species in the Ophioglossaceae. A timetable for gametophyte and young sporophyte development is postulated. Spores germinate soon after they are released in the spring, and mature gametophytes develop by the next growing season. Fertilization occurs approximately one year after spore dispersal, and after two years, the photosynthetic first leaf of the young sporophyte emerges.     

Nahrungserwerb und Konkurrenz des Blutspechts,Picoides (Dendrocopos) syriacus

Summary The Syrian Woodpecker, Picoides syriacus, acquires its food by excavating, probing, gleaning and fly-catching (Fig. 1). The food-acquisition is carried out from ground level to the crown layer of trees and beyond (Fig. 2).In an ecological sense the Syrian Woodpecker occupies an intermediate position between the Great Spotted and the Middle Spotted Woodpecker (Picoides major and medius). That is true for food-acquisition since it excavates more than medius and less than major whilst probing more than major and less than medius. The species is also intermediate in the choise of base from which foraging is done. Great Spotted Woodpeckers are found frequently on thin twigs and Middle Spotted Woodpeckers on thick stems.Between Great Spotted and Syrian Woodpecker there exist interspecific territoriality. Both are neutral with respect to the Middle Spotted Woodpecker. In spite of strong competition for nesting holes and frequent aggression the territories of Syrian Woodpecker and Starling (Sturnus vulgaris) overlap. These findings are discussed.     

Developmental morphology of strap-shaped gametophytes of Colysis decurrens: a new look at meristem development and function in fern gametophytes

Background and Aims

The gametophytes of most homosporous ferns are cordate–thalloid in shape. Some are strap- or ribbon-shaped and have been assumed to have evolved from terrestrial cordate shapes as an adaptation to epiphytic habitats. The aim of the present study was to clarify the morphological evolution of the strap-shaped gametophyte of microsoroids (Polypodiaceae) by precise analysis of their development.

Methods

Spores of Colysis decurrens collected in Kagoshima, Japan, were cultured and observed microscopically. Epi-illuminated micrographs of growing gametophytes were captured every 24 h, allowing analysis of the cell lineage of meristems. Light microscopy of resin-sections and scanning electron microscopy were also used.

Key Results

Contrary to previous assumptions that strap-shaped Colysis gametophytes have no organized meristem, three different types of meristems are formed during development: (1) apical-cell based – responsible for early growth; (2) marginal – further growth, including gametophyte branching; and (3) multicellular – formation of cushions with archegonia. The cushion is two or three layers thick and intermittent. The apical-cell and multicellular meristems are similar to those of cordate gametophytes of other ferns, but the marginal meristem is unique to the strap-shaped gametophyte of this fern.

Conclusions

The strap-shaped gametophytes of C. decurrens may have evolved from ancestors with a cordate shape by insertion of the marginal meristem phase between the first apical-cell-based meristem and subsequent multicellular meristem phases. Repeated retrieval of the marginal meristem at the multicellular meristem phase would result in indefinite prolongation of gametophyte growth, an ecological adaptation to epiphytic habitats.     

Arbuscular mycorrhiza formation in cordate gametophytes of two ferns, Angiopteris lygodiifolia and Osmunda japonica

Mycorrhizal symbiosis is common among land plants including pteridophytes (monilophytes and lycophytes). In pteridophytes with diplohaplontic life cycle, mycorrhizal formations were mostly reported for sporophytes, but very few for gametophytes. To clarify the mycorrhizal association of photosynthetic gametophytes, field-collected gametophytes of Angiopteris lygodiifolia (Marattiaceae, n = 52) and Osmunda japonica (Osmundaceae, n = 45) were examined using microscopic and molecular techniques. Collected gametophytes were mostly cut into two pieces. One piece was used for light and scanning microscopic observations, and the other for molecular identification of plant species (chloroplast rbcL sequences) and mycorrhizal fungi (small subunit rDNA sequences). Microscopic observations showed that 96 % (50/52) of Angiopteris and 95 % (41/43) of Osmunda gametophytes contained intracellular hyphae with arbuscules and/or vesicles and fungal colonization was limited to the inner tissue of the thick midribs (cushion). Fungal DNA analyses showed that 92 % (48/52) of Angiopteris and 92 % (35/38) of Osmunda have sequences of arbuscular mycorrhizal fungi, which were highly divergent but all belonged to Glomus group A. These results suggest that A. lygodiifolia and O. japonica gametophytes consistently form arbuscular mycorrhizae. Mycorrhizal formation in wild fern gametophytes, based on large-scale sampling with molecular identification of host plant species, was demonstrated for the first time.     

Temperature requirements for growth and maturation of the warm temperate kelp Eckloniopsis radicosa (Laminariales,Phaeophyta)

The annual kelp Eckloniopsis radicosa is distributed along Japanese coasts and occurs within the area with a February isotherm ranging 15–18°C and August isotherm ranging 25–28°C. In this study, the effects of temperature on the gametophyte growth and maturation, and the young sporophyte growth of E. radicosa were examined and the results are discussed in relation to the distribution of other warm‐adapted kelp species and the potential effects of climate change. The optimal temperature ranges for growth of male and female gametophytes were 23–27°C and 20–26°C, respectively. The upper survival temperature for gametophyte growth was 31°C for males and 30°C for females, respectively. The optimal temperature range for maturation of female gametophytes was ≤23°C. The optimal temperature range for growth of young sporophytes was 14–22°C. It was clarified that E. radicosa has the most warm‐adapted characteristics for growth and maturation of gametophytes among members of the Laminariales studied so far. The natural seawater temperature ranges during the growth and maturation seasons for gametophytes of E. radicosa, as well as the growth season for young sporophytes near to the northern and southern distribution limits (Izu‐Oshima: 14.9–24.5°C, Ichiki‐kushikino: 17.1–29.6°C), agreed with the experimentally determined temperature requirements. The warm‐adapted gametophyte stage and annual lifecycle are major factors enabling survival of E. radicosa in warm waters near tropical regions along the Japanese coast.     

PROBLEMS IN THE IDENTITY AND ORIGIN OF THE APPALACHIAN VITTARIA GAMETOPHYTE,A SPOROPHYTELESS FERN OF THE EASTERN UNITED STATES

Populations of a fern gametophyte presumed to be of the genus Vittaria occur commonly in the uplands of the southeastern United States. The gametophytes occur on non-calcareous rock outcrops of various composition in areas which provide continuous moisture and protection from temperature extremes. Gametophytes in these habitats are robust and long lived, frequently forming the dominant vegetation in areas covering several square feet. Reproduction is exclusively vegetative by production of gemmae. Although sex organs are present in most populations, viable sporophytes are never produced. The gametophytes have previously been considered most likely to be V. lineata, which occurs in Florida, but morphological and physiological comparisons do not support that conclusion. Significant differences between Appalachian and Florida gametophytes occur in growth form, growth rate, cold hardiness, sporophyte production, and patterns of gemma production. The distribution of the Appalachian gametophytes correlates with old, unglaciated land masses suggesting antiquity rather than recent introduction. Present evidence favors the interpretation that the Appalachian Vittaria gametophytes either belong to a tropical American species from which they have long been separated, or that they represent a distinct species of which the sporophyte no longer exists.     

A methodology for large-scale Athyrium sheareri gametophyte proliferation and sporophyte production using tissue culture

Tissue culture methods using gametophytes are considered the easiest ways to mass-produce fern sporophytes. The aim of this study was to develop a practical propagation method for the ornamental fern, Athyrium sheareri. The gametophytes obtained from in vitro spore germination were used as experimental materials. We used the chopping method to investigate the culturing conditions for proliferating gametophytes and the blending method for evaluating the mass production of sporophytes in mixed soil. Gametophyte proliferation was determined via Knop medium, various concentrations of Murashige and Skoog (MS) basal medium (1, 1/2, 1/4), and media components (sucrose, nitrogen source, and activated charcoal). The fresh weight of the gametophytes increased by more than 24-fold in 1/2 MS medium. In addition, 1 g of gametophyte could produce a maximum of 255.3 sporophytes in a mixed soil of 7.5 cm² area. Treating gametophytes with exogenous plant growth regulators promoted the formation and growth of sporophytes. The cultivated young sporophytes were acclimated and successfully grown in greenhouses. We developed a mass production protocol for A. sheareri sporophytes suitable for field application, which is expected to have commercial value.

     

Effect of temperature and photon fluence rate on gametophytes and young sporophytes of Laminaria ochroleuca Pylaie

We analysed the effects of temperature and photon fluence rate on meiospore germination, growth and fertility of gametophytes, and growth of young sporophytes of Laminaria ochroleuca. Maximum percentages of germination (91–98%) were obtained at 15°C and 18°C, independent of photon fluence rate. Optimal development of female gametophyte and maximum fecundity and reproductive success of gametophytes occurred at 15°C and 18°C and at 20 and 40 μmol m^–2 s^–1. Maximum relative growth rate of young sporophytes after 2 weeks of culture was achieved under the same conditions. L. ochroleuca gametophytes cannot reproduce and growth of its sporophytes is not competitive at temperatures close to 10°C. Received in revised form: 31 August 2001 Electronic Publication     

Arbuscular mycorrhizal colonization in field-collected terrestrial cordate gametophytes of pre-polypod leptosporangiate ferns (Osmundaceae,Gleicheniaceae, Plagiogyriaceae,Cyatheaceae)

To determine the mycorrhizal status of pteridophyte gametophytes in diverse taxa, the mycorrhizal colonization of wild gametophytes was investigated in terrestrial cordate gametophytes of pre-polypod leptosporangiate ferns, i.e., one species of Osmundaceae (Osmunda banksiifolia), two species of Gleicheniaceae (Diplopterygium glaucum, Dicranopteris linearis), and four species of Cyatheales including tree ferns (Plagiogyriaceae: Plagiogyria japonica, Plagiogyria euphlebia; Cyatheaceae: Cyathea podophylla, Cyathea lepifera). Microscopic observations revealed that 58 to 97 % of gametophytes in all species were colonized with arbuscular mycorrhizal (AM) fungi. Fungal colonization was limited to the multilayered midrib (cushion) tissue in all gametophytes examined. Molecular identification using fungal SSU rDNA sequences indicated that the AM fungi in gametophytes primarily belonged to the Glomeraceae, but also included the Claroideoglomeraceae, Gigasporaceae, Acaulosporaceae, and Archaeosporales. This study provides the first evidence for AM fungal colonization of wild gametophytes in the Plagiogyriaceae and Cyatheaceae. Taxonomically divergent photosynthetic gametophytes are similarly colonized by AM fungi, suggesting that mycorrhizal associations with AM fungi could widely occur in terrestrial pteridophyte gametophytes.     

Ontogenetic histology of Stegosaurus plates and spikes

Abstract: The dinosaur Stegosaurus is characterized by osteoderms of alternating plates and terminal paired spikes. Previous studies have described the histological features and possible functions of these osteoderms. However, ontogenetic changes are poorly documented. In this study, the ontogenetic changes of the osteoderms are examined using eight different ontogenetic skeletons (a juvenile, a subadult, a young adult, and five old adults based on the cortical histology of their body skeletons). The juvenile plate and subadult spike show thin cortex and thick cancellous bone. The young adult plates have an extensive vascular network, which is also seen in old adults. Old adult spikes are different from old adult plates in having a thick cortex and a large axial channel. The cortical histology, in both plates and spikes, show well‐vascularized bone tissue consisting of dense mineralized fibres in young adult forms. In old adult forms, the bone tissues in the spikes become more compact and are extensively remodelled. This might contribute to the structural reinforcement of the spikes. The plates in old adult forms also show extensive remodelling and lines of arrested growth, but only limited signs of compaction. The timing for acquisition of features seen in old adults is different between plates (an extensive vascular network in the young adult) and spikes (a thick cortex with a large axial channel in old adults). The result suggests that the timing for plate and spike functions is different. The extensive vascular networks seen in large plates suggest their function is for display and/or thermoregulation. The thick cortical bone of spikes of old adults suggests that spikes acquire a weapon function for defence ontogenetically late.     

CHARACTERIZATION OF MUCILAGE ON THE PROXIMAL CELLS OF YOUNG GAMETOPHYTES OF BOTRYCHIUM DISSECTUM (OPHIOGLOSSACEAE)

Young gametophytes of Botrychium dissectum produce a mucopolysaccharide coating on the external surface of the proximal cell. Scanning electron microscopy reveals deposition of the mucilage, which initially has a patchy distribution, in the shape of a thick triangular ring. Young gametophytes in the hydrated condition have a thin coating of mucilage over the other areas of the proximal cell wall. Histochemical staining indicates that the mucilage contains sugars with vicinal hydroxyl groups, carboxylated sugars, and small amounts of sulfated sugars. Protein and phenolic materials are also found in the mucilage. Lipids, β, 1-3 glucans and β,1-4 glucans are not present. Fluorescein labelled lectin binding shows the presence of terminal galactose and terminal fucose units. Germinating spores in the presence of the microtubule inhibitors CIPC (3-chloro-N-phenyl isporopyl carbamate) and griesiofulvin cause abnormal mucilage secretion. The inhibitor CIPC prevents mucilage secretion whereas griesiofulvin disrupts the pattern of deposition. Several functions of the mucilage are postulated.     

Macroscopic and histological investigation of guanaco footpads (Lama guanicoe,Müller 1776)

The surface of guanaco footpads is characterized by hairless skin with up to 4‐mm‐thick stratum corneum that protects from abrasion. The horny layer is pliable and elastic, and ensures firm contact with irregular ground. It is padded with a particular structure of the subcutaneous layer, the digital cushion. The flat cushions of each of the two digits are of elongated ovate shape, each about 45‐mm long, up to 20‐mm wide, and 8‐mm thick. The cushions are lined by a 1–2‐mm‐thick capsule that resembles a tunica albuginea. The capsule consists of coarse collagen fibers, with elastic fibers absent. The cushion capsule and dermis approach each other, and fuse along a line that runs parallel to the longitudinal axes of cushion and digit. Loose connective tissue rich in elastic fibers and acidic glycosaminoglycans separates dermis and cushion capsule lateral to the narrow interconnecting zone. The cushion capsule encloses cloudy yellowish, gelatinous material. Microscopy shows bundles of elastic fibers in abundant mucinous matrix. Tightly gathered elastic bundles adjoin the inner surface of the capsule. Rough cords of elastic fibers branch out from there and traverse to the opposite side. The cushion is pressed flat, and elastic fibers are stretched when bearing weight. With relief of load, elastic fibers contract and reset the cushion's shape. Contractile cells are absent. A resistant capsule and easily malleable mucinous contents establish the functioning as a gel pad. Mucinous connective tissue between elastic fiber bundles contains abundant basophilic matrix. Hyaluronan, chondroitin sulfate, and dermatan sulfate are main matrix constituents. Spindle‐shaped or stellate fibroblasts contain vimentin, S100 protein, and neuron specific enolase. Moprhology, staining characteristics and synthesis activities of these cells meet the criteria to be classified as myxoid cells. The connective tissue in guanaco digital cushions represents myxoid tissue. J. Morphol. 276:331–341, 2015. © 2014 Wiley Periodicals, Inc.     

The effects of light on sex determination in gametophytes of the fern <Emphasis Type="Italic">Ceratopteris richardii</Emphasis>

The sexuality of homosporous fern gametophytes is usually determined by antheridiogen, a pheromone that promotes maleness. In this work the effect of photomorphogenically active light on antheridiogen-induced male development was examined for gametophytes of Ceratopteris richardii. Although blue light did not affect sensitivity to Ceratopteris antheridiogen (A_Ce) in wild-type gametophytes, it was found that the gametophytes of the her1 mutant, which are insensitive to A_Ce, developed into males when grown under blue light in the presence of A_Ce. Thus, we conclude that another A_Ce-signal transduction pathway activated by blue light exists latently in the gametophytes of C. richardii. Red light, on the other hand, suppressed male development. Because simultaneous red and blue light-irradiation did not promote male development in the her1 gametophytes, the action of red light seems to dominate that of blue light. The results of experiments with a photomorphogenic mutant also suggested that phytochrome may be involved in the action of red light.     

环境因子对中华双扇蕨孢子萌发的影响

为了解珍稀濒危植物中华双扇蕨(Dipteris chinensis)濒危原因,研究了温度和湿度对其孢子萌发的影响.结果表明,中华双扇蕨孢子萌发能力较强,萌发方式为Vittaria型,配子体萌发为Marattia型.中华双扇蕨孢子繁殖不受温度影响,而湿度显著影响孢子繁殖过程,湿润环境中孢子正常萌发,并形成心形配子体,顺利...     

Gametophyte contribution to sporophyte growth on the basis of carbon gain in the fern <Emphasis Type="Italic">Thelypteris palustris</Emphasis>: effect of gametophyte organic-matter production on sporophytes

At an early stage of growth gametophytes support the sporophytes of ferns. Young sporophytes become independent of gametophytes when the first leaves develop. Although large fern gametophytes produce multiple archegonia simultaneously, only one sporophyte is typically established on one gametophyte. The number of sporophytes is believed to be controlled in two possible directions, from gametophyte to sporophyte or from preceding sporophyte to another sporophyte. To investigate the effects of gametophytes on their sporophytes, we studied the relationship between organic matter production by gametophytes and the growth of young sporophytes of Thelypteris palustris. We cut gametophytes in half (CGs) to reduce the gametophytes’ production of matter. There was no significant difference between the growth of sporophytes on intact gametophytes (IGs) and that on CGs. According to our estimates, based on the rate of organic matter production, the large gametophyte was able to produce two or more sporophytes. The resources required for CGs to make similar-sized sporophytes was twice that for IGs. In polyembryony each of the multiple sporophytes was similar in size to the single sporophytes. Resource limitation does not seem to explain why fern gametophytes establish single sporophytes.     

Contribution of fern gametophytes to the growth of produced sporophytes on the basis of carbon gain

Sporophytes appeared on most gametophytes of Thelypteris palustris (Salisb.) Schott that reached a certain size, which is interpreted to be a critical size of gametophytes for the production of sporophytes. After sporophytes were produced, attached gametophytes ceased dry weight growth, but the gametophytes which did not produce sporophytes grew successively. It was hypothesized that matter produced by gametophytes was being supplied to young sporophytes. Photosynthesis and respiration of gametophytes with attached sporophytes were not significantly different from that of gametophytes without sporophytes. Photosynthetic activity of gametophytes dropped from 0.18 to 0.03 mol CO₂ g^–1 s^–1 during the growth period. The higher photosynthetic rates of gametophytes in the early growth stage were important for reaching the critical size for sporophyte production in a short time. Sporophytes in the one leaf stage averaged 0.14 mol CO₂ g^–1 s^–1 of photosynthetic activity. The results show that sporophytes that had expanded the first leaf grow by their own photosynthetic production. Gametophytes allocated the photosynthate for sporophytes and it was an important aid before the one-leaf stage. The supportive role of gametophytes ended at that stage.     

THE GAMETOPHYTES OF OPHIOGLOSSUM PALMATUM L.

Gametophytes of the epiphytic species Ophioglossum palmatum L. are described for the first time, bringing the number of Ophioglossum species with known gametophytes to 11. Although somewhat unusual in terms of (a) degree of branching, (b) the marked expansion of the basically cylindrical axis in some cases, and (c) production of stout, papilla-like rhizoids, the gametophytes of O. palmatum are more similar to other Ophioglossum gametophytes than they are to the gametophytes of Botrychium or Helminthostachys. Gametophyte characters do not provide strong evidence for placing O. palmatum into a separate genus, viz., Cheiroglossa. The gametophytes of O. palmatum most closely resemble those of the other epiphytic species in the genus, O. pendulum. This similarity suggests a need for a revaluation of modern taxonomies which, in general, treat the two species as only distantly related.