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morphology of ferns

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July 8, 2013

morphology of ferns

WANG Guo-Hong, FANG Jing-Yun, GUO Ke, XIE Zong-Qiang, TANG Zhi-Yao, SHEN Ze-Hao, WANG Ren-Qing, WANG Xiang-Ping, WANG De-Li, QIANG Sheng, YU Dan, PENG Shao-Lin, DA Liang-Jun, LIU Qing, LIANG Cun-Zhu. Lond. Cutter, E. G. (1978). Furthermore, ferns grow in many habitats—from mangroves at sea level to alpine vegetation above tree line, temperate forests to arctic tundras, and desserts to wetlands. The sterile leaves of Psilotum (Psilotaceae) are unusual among ferns by being scale-like and generally lacking a vascular supply. Faculty Sci. The second reason for the uncertainty about megaphyll homology in ferns is that there are conflicts about the interpretation and codification of characters of extinct Devonian and Carboniferous fernlike plants without laminate leaves. Besides the typical characteristics of all megaphylls, fern leaves have some additional characteristics that make them distinctive. Two genera of ferns that have indeterminate leaves twining around a support are Salpichlaena (Blechnaceae) and Lygodium (Lygodiaceae, Figure 2H; Mueller, 1983a,b). The leaves of rheophytic ferns exhibit several morphological and anatomical adaptations to their unusual habitat. (N) Anemia adiantifolia, hemidimorphic, with two basal pinnae fertile and long-stalked. Sec. In addition, experimental studies found that fern leaves are not determined as leaves until later in their development but when first specified have more of a shoot identity (reviewed in White, 1971). doi: 10.1007/BF02489484, Imaichi, R. (1982). Phytomorphology 11, 346–359. 95, 435–453. 12, 617–628. A colorless ventral lobe rests on the water and a thicker green dorsal lobe arches upward. doi: 10.5962/bhl.title.26129. A linear sequence of extant families and genera of lycophytes and ferns. No use, distribution or reproduction is permitted which does not comply with these terms. Generally such leaves are either pendulous or scrambling over the surrounding vegetation. Toward a new circumscription of the twinsorus-fern genus (. Bot. 64, 125–152. A community-derived classification for extant lycophytes and ferns. In vitro morphogenesis in Osmunda cinnamomea. Van den Heede CJ, Viane RLL, Chase MW ( 2003). This hypothesis is based on the statement that the earliest known megaphyll-like structures are highly dissected and composed of segments that were short, narrow, and single-veined, but lacked an expanded lamina and the abaxial/adaxial anatomical organization of leaves (Rothwell, 1999; Boyce and Knoll, 2002). Fern leaves and megaphylls of other groups are defined by a combination of characters that are a result of specific developmental processes. In all vascular plants, leaves arise from the SAM. These conflicts make the phylogenetic placement of the fossils equivocal, and therefore statements of leaf homology within ferns ambiguous. J. These three processes transformed the dichotomous, leafless, photosynthetic axes of the sporophyte of early Devonian plants into leaves (Zimmermann, 1930, 1952; Wilson, 1953; Stewart, 1964). A new development: evolving concepts in leaf ontogeny. Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Zhang GM, Liao WB, Ding MY, Lin YX, Wu ZH, Zhang XC, Dong SY, Prado J, Gilbert MG, Yatskievych G, Ranker TA, Hoope EA, Alverson ER, Metzgar JS, Funston AM, Masuyama S, Kato M ( 2013). Hypotheses of megaphyll homology depend on the morphological and anatomical interpretations of extinct leafless Devonian and Carboniferous plants. Given this great diversity in functions and habitats it is not surprising that fern leaves exhibit a great diversity in size and shape (Figures 2, 3). In other ferns, the leaf bases store abundant starch and are termed trophopods (Wagner and Johnson, 1983). doi: 10.1086/330590. Am. Fahn, A. Their laminae often exhibit constrictions where fiddlehead activity diminishes during a less favorable part of the growing season. $2.50 The Power of Movement in Plants. doi: 10.1242/dev.033811, Kuehnert, C. C. (1967). Science 296, 1858–1860. The dimorphy may be of two types. Fern Structure Leaves. doi: 10.2307/2418896. The phylogenetic position and delimitation of Pteridaceae were treated differently by different authors. The occurrence of intercalary and uninterrupted meristems in the internodes of tropical monocotyledons. There has been recent interest in extending leaf evolutionary developmental studies to other species and lineages, particularly in lycophytes and ferns. In contrast, seed plants (with few exceptions) have chloroplasts only in the guard cells of the epidermis, not in other epidermal cells (Cutter, 1978; Gifford and Foster, 1988; Fahn, 1990). doi: 10.1086/600135, Sano, R., Juárez, C. M., Hass, B., Sakakibara, K., Ito, M., Banks, J. Development 138, 2925–2934. 63, 852–856. FENG Si-Yuan, WEI Ya-Nan, WANG Zhen-Juan, YU Xin-Yang. The number of times leaves are thought to have evolved separately in vascular plants depends on the phylogenetic hypothesis used and the inclusion and morphological interpretations of fossil taxa (Boyce, 2010). The Salvinia paradox: superhydrophobic surfaces with hydrophilic Ppins for air retention under water. Morphology of some polystichoid ferns Morphology of some polystichoid ferns CHANDRA, PRAKASH; NAYAR, B. K. 1970-07-01 00:00:00 The morphology of the spores and prothalli of Arachniodes aristata, A. assamica, Cyrtomium caryotideum, C. falcatum and 10 species of Polystichum is described. ; Polypodiaceae), and Vittaria (6 spp. 169-256. Experiments on the cause of dorsiventrality in leaves. TANG Li-Li,ZHANG Mei,ZHAO Xiang-Lin,KANG Mu-Yi,LIU Hong-Yan,GAO Xian-Ming,YANG Tong,ZHENG Pu-Fan,SHI Fu-Chen. A comparison of the basal scales and Indusia of, Holttum ER ( 1957). 19, 417–437. Philos. Pteridophytes and Gymnosperms Vol. Developmental potentialities of leaf primordia of Osmunda cinnamomea. These structures occur in Cystopteris protrusa (Cystopteridaceae) and Onoclea sensibilis (Onocleaceae). Can. Morphology of Vascular Plants. However, if incisions were made between the SAM and leaf primordia at developmental stage P4, then these isolated primordia were more likely to grow out as leaves (Cutter, 1954, 1956). Key morphological alterations in the evolution of leaves. A., Hasebe, M., and Juarez, C. M. (2005). Leaf primordia P1, P2, and P3 are more plastic in their development and after incisions, may grow out as buds, although a few still develop as leaves. Some fern leaves present unusual shapes and adaptations. The morphology of pteridophytes; the structure of ferns and allied plants Item Preview remove-circle Share or Embed This Item. London: MacMillan and Co. Boyce, C. K. (2010). Lobed leaves are also found in ferns, and the basic plan may be either pinnate, as in Aglaomorpha meyeniana (Polypodiaceae, Figure 3Y), or palmately lobed as in Hemionitis palmata (Pteridaceae, Figure 2G). Lond. In these cases, fertile leaves may be produced at certain times of the year, such as during the wet season or dry season. Plant Biol. 3(Suppl. Wagner, W. H. Jr., and Johnson, D. M. (1983). An interpretation of the morphology and evolution of the cone and shoot of Equisetum. For example, developmental genetic studies in ferns with diverse morphologies (e.g., simple vs. compound leaves) could provide the molecular basis for their morphological diversity. doi: 10.1666/0094-8373(2002)028<0070:EODPAT>2.0.CO;2. Corvez, A., Barriel, V., and Dubuisson, J.-Y. Greenfield SS ( 1938). Ferns and Allied Plants. doi: 10.1038/164167a0. The inclusion of Psilotaceae within the ferns suggests that the small scale-like appendages in Psilotum are probably best interpreted as highly reduced leaves. At least in fossil ferns, the defined adaxial/abaxial identities are also present in the vascular bundles of the petiole where the protoxylem is adaxial (in contrast to seed plants that have abaxial protoxylem; Galtier, 2010). Ferns and fern relatives are known as Pteridophytes. Most people probably envision ferns this way because, in fact, most fern leaves are highly divided. Philipp. 91, 717–727. Plant Cell 22, 1019–1032. The results from experimental leaf studies performed in ferns parallel the results found in angiosperms. Presumably, the function of coiling is to protect the soft meristematic parts concealed within the fiddlehead. This is not a thigmotropic grasping response as exhibited, for instance, by the leaves of Clematis (Ranunculaceae; Darwin, 1876). J. Bot. 171, 641–661. Rev. 63, 510–525. Another hypothesis proposes that megaphylls in the euphyllophytes may be homologous at the level of lateral branches (megaphyll precursors), and that the processes of flattening into one plane (planation, which sometimes also has been suggested to imply abaxial/adaxial anatomical organization of leaves) and the formation of a lamina (webbing) developed independently in ferns and seed plants (Kenrick and Crane, 1997; Galtier, 2010). (Q) Gleichenia microphylla, pair of opposite pinnae. Series [Hutchinson university library] Biological sciences, Hutchinson university library. They are arranged in a fixed and predictive phyllotaxy around the shoot apex (Schoute, 1938; Gifford and Foster, 1988), they have adaxial/abaxial identities and with laminar tissue supplied by numerous vascular traces, and most grow for a finite amount of time to a finite size (Bower, 1923; Hagemann, 1989; Kaplan and Groff, 1995; Kaplan, 2001). B 175, 565–615. Morphogenetic studies on Osmunda cinnamomea L. The origin and early development of vegetative fronds. 1. Gifford, E. M., and Foster, A. S. (1988). Distal to this, the laterally expanded portion of the leaf is termed the blade or lamina, whose central midrib is referred to as the rachis. Presentedby HillaryHouse PublishersLtd, Jan.8,1962 s^^^^^s These studies found that, as in seed plants with compound leaves, Class I KNOX genes are expressed in the meristem and in the developing leaves. Received: 28 May 2013; Accepted: 15 August 2013; Published online: 04 September 2013. Experimental induction of buds from fern leaf primordia. doi: 10.2307/1309213, Nagalingum, N. S., Schneider, H., and Pryer, K. M. (2006). Can. Is Ophioglossum palmatum anomalous. The specification of adaxial and abaxial identities has been suggested to be important for lamina outgrowth (Waites and Hudson, 1995). of fern morphology is the presence of epidermal ap-pendages. Yet unlike seed plants, Class I KNOX genes were not found to be down-regulated in leaf primordia. Watkins Jr JE, Kawahara AY, Leicht TA, Auld JR, Bicksler AJ, Kaiser K ( 2006). Hicks, G. S., and Steeves, T. A. doi: 10.1007/BF02868687, White, R., and Turner, M. (1995). The gametophytes of many epiphytic ferns consequently have a much more branched and dissected morphology than their terrestrial counterparts (either ribbon-shaped, filamentous, or strap-shaped), which is capable of continued meristematic growth (Figure 7) (Dassler and Farrar, 2001). Plant Sci. Further studies on the influence of determined leaf primordia on undetermined leaf primordia. Evolut. This prolonged meristematic activity is due to an apical cell at the leaf tip and is a distinctive feature of ferns compared to seed plants (Imaichi, 2008). (Z) Cyrtomium macrophyllum, 1-pinnate. In those few fern species that invest their fiddleheads in a thick covering of mucilage, the lines are modified at the pinna bases into elongate peg-like structures that protrude through the mucilage, thus aerating the developing leaf (Hennipman, 1968). Can. Some plants that are called ferns, such as asparagus ferns, reproduce by seeds and are not true ferns. Can. White, R. (1971). Curr. Darwin, C. (1896). Bot. Rothwell, G. W. (1999). Plant Physiology. These lines aerate the leaf. As an adaptation to their epiphytic life style, the drynarioid genera of Polypodiaceae have leaves modified for collecting organic debris that falls from above, mostly bits of bark and leaves (Hennipman and Roos, 1982; Janssen and Schneider, 2005). Int. 47, 59–63. Sector analysis and predictive modelling reveal iterative shoot-like development in fern fronds. J. Bot. There is usually a stalk (the stipe) with a flat blade (the lamina), often divided into segments. Although primarily for photosynthesis, fern leaves may also assume other tasks such as propagating the plant vegetatively by bulblets, harboring nitrogen-fixing cyanobacteria, forming nests that collect humus falling from above, or efficiently dispersing spores. doi: 10.1007/BF02488625, Janssen, T., and Schneider, H. (2005). Most studies use the terminology “P” and “I” where P1 is the youngest leaf primordium and P10 is the 10th oldest primordium. In other ferns, such as Bolbitis heteroclita (Dryopteridaceae, Figure 2O), Thelypteris reptans (Thelypteridaceae, Figure 2N) or Asplenium rhizophyllum (Aspleniaceae), the lamina apices are long-attenuate or flageliform (whip-like) with buds along their length or at their tips. (A) Eupodium laeve (Marattiaceae). Mesquite: a modular system for evolutionary analysis. From genes to shape: regulatory interactions in leaf development. 123, 43–55. (1907). Ferns are the most diverse group of vascular plants after seed plants. The lack of these fossils means that there is no evidence concerning the mode of origin of their leaves. In support of their results in lycophytes, Harrison et al. (1990). Epidermal outgro wth often takes the form. Features of scales play an important role in classification of ferns. 232, 343–384. Maturation of the developing fern leaf is acroscopic, that is, toward the apex (Figure 5). (U) Hemionitis ariifolia, hastate at left, deltate at right (from same plant). Developmental potentialities of leaf primordia of Osmunda cinnamomea. Horsetails (Equisetopsida) and whisk ferns (Psilotales) are treated as part of the fern lineage. A rhizome is a specialized, root-like stem. doi: 10.1016/j.pbi.2007.07.012, Pubmed Abstract | Pubmed Full Text | CrossRef Full Text, Barthlott, W., Schimmel, T., Wiersch, S., Koch, K., Brede, M., Barczewski, M., et al. Sharpe, J. M., and Mehltreter, K. (2010). No_Favorite. The effect of various physiologically active substances on the development of marsilea in sterile culture. (1948b). Development 121, 2143–2154. Its blade-less leaf is interpreted as a petiole that has lost its apical pinnae (Eames, 1936). Morphology and Anatomy. 55, 223–231. V. Toward greater understanding of the final morphogenetic expression of isolated set I cinnamon fern leaf primordia. Microsurgery and sterile culture experiments demonstrated that the SAM does not require attached leaf primorida or leaves to continue the development of the adult shoot and therefore the SAM is capable of autonomous development. Philos. A conspectus of the native and naturalized species of Nephrolepis (Nephrolepidaceae) in the World. A monograph of the fern genus Eriosorus. Isolated P3 primordia were grown on media containing homogenized leaves (P10 or P12), which resulted in most of P3 primordia developing as leaves (Kuehnert, 1969b; White, 1971). Narrower laminae or pinnae have a thinner boundary layer of air (Salisbury and Ross, 1992). J. Bot. Jiuxiang Huang,Wenna Chen,Yuling Li,Gang Yao. Studies in ferns are important for resolving morphological interpretations within particular fern groups and are crucial to our understanding of leaf evolution and development in vascular plants. 22, 2325–2328. Phyllotaxis. Lond. The other kind of leaf is whitish, highly divided, and hanging down in the water. doi: 10.1146/annurev.ecolsys.29.1.567, Doyle, J. A KNOX family TALE. R. Soc. (1910). Being the main conspicuous organ of nearly all vascular plants and often easy to recognize as such, it seems surprising that leaves have had multiple origins. 11, 45–67. This is important nutritionally because these ferns are epiphytic, not in contact with water and minerals in the soil. Wei R, Schneider H, Zhang XC ( 2013). The phylogenetic relationship of Ophioglossaceae. (H) Lygodium flexuosum, rachis (at right) with lamina of pinnule (other half of pinna not shown). Initiation of the first and the second petiolar buds in relation to early leaf ontogeny. Examples include certain species of Gleicheniaceae (Figures 2Q, 8; Moran, 2004), Jamesonia (Pteridaceae, Tryon, 1970), and Hypolepis (Dennstaedtiaceae, Brownsey, 1987; Holtum, 1958). Tokyo Sec. 47, 65–68. Sci. (F) Actiniopteris semiflabellata, incised leaf. Although typically envisioned as compound, the leaves of ferns actually display great morphological diversity (Figures 2, 3). The rhizome of Oleandra pistillaris and 0. wallichii is elongated (erect in the former, creeping in the latter), cylindrical, clothed with peltate, non-glandular, hairy paleae and bearing leaves in small clusters separated by long leafless portions. The humus-collecting leaves are also brown, stiff, papery, and dead at functional maturity. doi: 10.1139/b67-229, Kuehnert, C. C. (1969a). 46, 957–968. They develop from cell divisions of several epidermal cells. False veins occur in some filmy fern genera, such as Didymoglossum (Wessels Boer, 1962). doi: 10.1600/036364409789271209. Chrysler, M. A. Developmental study on Hypolepis punctata (Thunb.) Sterile-fertile leaf dimorphy and evolution of soral types in Polybotrya (Dryopteridaceae). doi: 10.1111/j.1095-8339.1972.tb02279.x, Prigge, M. J., and Clark, S. E. (2006). The morphology of pteridophytes;: The structure of ferns and allied plants ([Hutchinson university library] Biological sciences): Books - For decades, morphologists, anatomists, paleobotanists, and systematists have contributed data to this debate. 91, 1726–1741. One hypothesis suggests that megaphylls are not homologous within the euphyllophytes. Proc. doi: 10.1111/j.1095-8339.1990.tb00909.x. Lond. In addition, isolated SAMs grown in sterile culture on media supplemented with sucrose and auxin formed adult plants (White, 1971). Buy the Paperback Book The Morphology of Pteridophytes; the Structure of Ferns and Allied Plants by K R Sporne at, Canada's largest bookstore. pp. Genera Hymenophyllacearum. New York, NY: Macmillan. doi: 10.1139/b92-022. doi: 10.2307/2441679, Moran, R. C. (1986). In some ferns the aerophores extend distally into the rachis, or proximally onto the rhizomes (e.g., Mickelia and Polybotrya, both Dryopteridaceae). A monograph of the fern genus Platycerium (Polypodiaceae). Dimorphism is a syndrome of many characters, and these may be anatomical or morphological. As the debris decomposes, it forms humus into which the plant grows roots to absorb water and nutrients. Is morphology really at odds with molecules in estimating fern phylogeny. These studies were designed to understand how fern leaves, and ferns in general, develop. 47, 481–488. Most ferns have specialized stems called rhizomes that are. The science of plant morphology: definition, history, and role in modern biology. Because ferns occupy a key phylogenetic position as sister to the seed plants (Figure 1), comparative studies in diverse fern species may help to elucidate the evolution of these leaf developmental regulators and their role in leaf development. J. Bot. There are two characteristics typical of most fern leaves: a fiddlehead and aerophore lines (Figure 6). Bot. Plant Anatomy. doi: 10.1139/b69-063, Harrison, C. J., Corley, S. B., Moylan, E. C., Alexander, D. L., Scotland, R. W., and Langdale, J. Opin. Schuettpelz E, Schneider H, Huiet L, Windham MD, Pryer KM ( 2007). Developmental potentialities of leaf primordia of Osmunda cinnamomea. Although some results showed that if I1 is isolated then it more likely develops as a bud, but in rare cases it developed as a leaf (Amer and Williams, 1957). Christenhusz MJM, Zhang XC, Schneider H ( 2011). Pryer KM, Schuettpelz E, Wolf PG, Schneider H, Smith AR, Cranfill R ( 2004). Sometimes the basal pinnae are repeatedly branched on the basiscopic side—a condition known as pedate. Ferns are seedless vascular plants of humid tropics and temperate areas. The root system is matted and tightly anchored to the substrate. (H) Salvinia molesta, root-like (lower) leaf is submerged and bears sori, the two round ones are floating. Evolution of the class III HD-Zip gene family in land plants. The thin laminae of filmy ferns dry out readily and then, upon rehydration, rapidly expand and resume photosynthesis (Proctor, 2003, 2012). More recently, molecular genetic studies have provided insight into leaf evolution and development mainly within angiosperms and, to a lesser extent, lycophytes. A glossary of some terms relating to the fern leaf. Cutter, E. G. (1956). Tweede Reeks 80, 1–126. Many of the experimental studies on fern leaf development that we will discuss were performed on two different leptosporangiate fern species Dryopteris aristata (Dryopteridaceae) and Osmunda cinnamomea (Osmundaceae). Schuettpelz, E., Schneider, H., Huiet, L., Windham, M. D., and Pryer, K. M. (2007). Ostrich Fern. (V) Vittaria lineata, linear leaves (shoe-string fern). (1989). Homologies in leaf form inferred from KNOXI gene expression during development. Tsutsumi C, Kato M ( 2008). Also, in almost all tropical ferns with strong sterile-fertile leaf dimorphy, the fertile leaves tend to be shorter-lived than the sterile. Here the petiole continues into the lamina as a single, unbranched rachis that produces lateral pinnae. Although primarily for photosynthesis, fern leaves may also assume other tasks such as propagating the plant vegetatively by bulblets, harboring nitrogen-fixing cyanobacteria, forming nests that collect humus falling from above, or efficiently dispersing spores. This is especially unknown for tropical ferns not limited by an unfavorable winter growing season and thus capable of producing leaves throughout the year. 2. Of great importance in fern taxonomy is the distinction between Eupolypods I and II (i.e., about 65% of extant ferns) based on petiole vasculature. (M) Matteuccia struthiopteris, sterile-fertile leaf dimorphism (fertile leaf at right). FANG Jing-Yun, GUO Ke, WANG Guo-Hong, TANG Zhi-Yao, XIE Zong-Qiang, SHEN Ze-Hao, WANG Ren-Qing, QIANG Sheng, LIANG Cun-Zhu, DA Liang-Jun, YU Dan. Rep. Tokyo Bunrika Daigaku B 15, 1–21. Bower, F. O. J. (1960). Instead of the stems common to most plants, ferns have rhizomes. Qi XP, Kuo LY, Guo CC, Li H, Li ZY, Qi J, Wang LB, Hu Y, Xiang JY, Zhang CF, Guo J, Huang CH, Ma H ( 2018). (P) Pellaea cordifolia, decompound. The absence of chloroplasts from the epidermis of seed plants (except the guard cells) is best interpreted on the basis of outgroup comparison as a loss, and this loss represents a synapomorphy for seed plants. “Morphology,” in Manual of Pteridology, ed F. Verdoorn (Hague: Martinus Nijhoff), 1–104. Molecular phylogenetic hypotheses of extant taxa consider ferns as a monophyletic group that includes the Equisetaceae, Psilotaceae, and Ophioglossaceae (Figure 1; Pryer et al., 2001; Qiu et al., 2006; Grewe et al., 2013). 2. Independent recruitment of a conserved developmental mechanism during leaf evolution. Rheophytes of the World. If the molecular phylogenetic hypothesis is accepted, then all ferns leaves should be homologous at some level. BMC Evolut. J. Bot. J. Bot. Briggs, W. R., and Steeves, T. A. (A) Lindsaea cyclophylla (Lindsaeaceae). Science 225, 1697–1699. Taxon 32, 268–269. doi: 10.1073/pnas.0603335103, Raubeson, L., and Jansen, R. (1992). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. One process involved the elongation of some of the branches more than others, producing a main central branch with subordinate lateral ones (overtopping). Articles. Even within ferns the homology of leaves is unclear. A phylogenetic classification of the homosporous ferns. A few ferns, however, have indeterminate leaves. Kenrick, P., and Crane, P. R. (1997). Sterile-fertile leaf dimorphy is common in ferns. Charact. 16, 1911–1917. Ferns have 3 major parts the rhizome, the fronds and the reproductive structures called sporangia. (1983b). Recent morphological and molecular phylogenetic analyses indicate that ferns are the sister group of seed plants (Raubeson and Jansen, 1992; Stevenson and Loconte, 1996; Kenrick and Crane, 1997; Pryer et al., 2001), and include the families Psilotaceae, and Equisetaceae, which have not always been considered as ferns (Figure 1; Pryer et al., 2001). The comparative ecology of San Ramon Polypodiaceae. Expression-potential of undetermined primordia separated by a barrier membrane from undetermined or determined primordia. Ferns are flowerless green plants . doi: 10.1093/pcp/pcs074. Philos. Marsilea (Marsileaceae, Figure 2K) is unusual because its leaves resemble a four-leaved clover. Ann. This thinner layer of air would promote drying of sporangia and increase the chance that spores, after being catapulted from sporangia, would soon encounter moving air currents. This thicker lobe contains a cavity that harbors nitrogen-fixing cyanobacteria (Anabaena azollae). London: E. Arnold. “Developmental themes in vascular plants: functional and evolutionary significance,” in Experimental and Molecular Approaches to Plant Biosystematics, eds P. C. Hoch and A. J. Stephenson (St. Louis, MO: Missouri Botanical Garden). doi: 10.1139/b69-009, Kuehnert, C. C. (1969b). More than 30 species in the collection are British Columbia natives. A molecular phylogeny of the fern family Pteridaceae: Assessing overall relationships and the affinities of previously unsampled genera. Holtum, R. E. (1958). Figure 8. Even though there is no complete sequenced genome or functional model system for ferns, there is now a considerable amount of transcriptome data for ferns available through the 1 KP project ( In contrast, the fertile leaves are green, much longer, and arching away from the substrate. The “simple” structure of the leaves and the dichotomizing axes that constitute the entire plant, led several authors to relate Psilotum to the earliest vascular plants (Bower, 1935; Eames, 1936; Wilson, 1953; Rothwell, 1999). Yet fern leaves exhibit enormous diversity, especially in size, shape, and cutting (Figures 2, 3). (C) Adiantum pedatum, pedate. (K) Davallia heterophylla, holodimorphic, fertile leaf at left. In addition, if I1 is isolated from the apex then the incipient primordium develops as a shoot. Rev. “Ophioglossaceae,” in The Families and Genera of Vascular Plants. (S) Astrolepis sinuata, 1-pinnate-pinnatifid. These represent 26 separate genera in 13 families. Figure 2. Comparative studies of the Class I KNOX genes have been performed in the leptosporangiate ferns Anogramma chaerophylla (Pteridaceae), Ceratopteris richardii (Pteridaceae), and Osmunda regalis (Osmundaceae, Figure 3O), all of which have divided leaves (Bharathan et al., 2002; Harrison et al., 2005; Sano et al., 2005). In addition, the signal from developing leaves involves a diffusible substance. In Equisetum, the relationship of the vegetative and fertile portions has been debated: the sporangiophore (the fertile portion) has been interpreted as being a novel organ “(organ sui generis)” or homologous to the leaves (Goebel, 1905; Bower, 1935; Zimmermann, 1952; Page, 1972). Byrne, M. (2012). doi: 10.2307/2443122, Mueller, R. J. Leaves are lateral determinate structures formed in a predictable sequence (phyllotaxy) on the flanks of an indeterminate shoot apical meristem. pp. Molecular genetic studies in angiosperm leaf development are beginning to provide clues into the nature of the signaling that occurs between the shoot apex and leaf primordia (Byrne, 2012). Can. The ground tissue of the rhizome is composed of thick-walled parenchyma, and there is a peripheral sclerenchymatous sheath as well as … Bot. Mét. 44, 1172–1185. Experimental and analytical studies of Pteridophytes: XVII. Blumea 50, 279–322. Trans. They bear stomata on their abaxial surfaces and have compact mesophyll with little intercellular spaces (Goebel, 1905). Natl. ; Pteridaceae). doi: 10.1007/s00606-004-0264-6, Kaplan, D. (2001). These pinnae, called aphlebiae, are of unknown function. Making leaves. Scales can be persistent in mature leaves and may become smaller and reduced to uniseriate proscales toward the margins of the laminae (Moran, 1986). (R) Adiantum raddianum, decompound. The first kind of leaf, of which there are two in the false whorl, is floating, green, entire, and conduplicate in bud (i.e., not circinate). One is conflicting phylogenetic hypotheses. Clearly these growth form characters have been derived independently in different epiphytic lines and reflect habitat selection rather than genetic ancestry. Similar results were found in Osmunda cinnamomea leaf primordia at developmental stages P1-P10 that were isolated and grown in sterile culture (Steeves, 1961, 1963; White, 1971). 47, 69–72. Opin. Ferns once dominated the earth in carboniferous period (about […] doi: 10.1007/BF00708854. Fiddleheads are highly distinctive of ferns because they are absent from lycophytes and nearly all seed plants (some exceptions in seed plants are the cycads Cycas and Stangeria, and the insectivorous plant Drosophyllum lusitanicum, although in the latter the coiling is abaxial, not adaxial as in ferns). Christenhusz MJM, Chase MW ( 2014). However, Wardlaw (1949b,d) concluded that initially leaf primordia and shoot buds of Dryopteris aristata are histologically indistinguishable. Also maximizing spore dispersal are the longer and more erect petioles that elevate the fertile lamina away from the substrate where the spores are more likely to be picked up by air currents. 600 spp. Fern laminar scales protect against photoinhibition from excess light. The leaf bases are sessile and wide so that the accumulated humus does not fall through. Among ferns, the few exceptions that lack chloroplasts in all epidermal cells are species that grow in full sun, such as high-canopy epiphytes (e.g., Elaphoglossum lingua, Dryopteridaceae) or on sunny rock faces (e.g., Notholaena affinis, Pteridaceae; Moran, pers. ”Ç©¶ 21, 360-364. 于顺利, æž—å°¤å ´ ( 1996 ) leaf.! Role in modern biology French, J. L. ( 1985 ) 2K ) is unusual its... < 0070: EODPAT > 2.0.CO ; 2 are generally considered to be important for lamina outgrowth ( and! Paradox: superhydrophobic surfaces with hydrophilic Ppins for air retention under water most people probably envision ferns this way,... The role of the leaf bases are sessile with deeply pinnatifid laminae and expanded bases turn! Been shown to play an important role in the collection are British Columbia natives present are. Plants that are divided multiple time along the length of the summer and therefore is no evidence concerning the of! Water relations and desiccation tolerance of the effects of homogenized, determined leaf primordia on of... Polycyclic condition found in Acrostichum ( Pteridaceae ) and N. pendula, exhibit pendulous indeterminate leaves (,... Two genes have been derived independently in different groups of vascular plants has widely. Biological sciences, Hutchinson university library and undivided or it may be homologous different. Diversity in functions and habitats it is not surprising that they have adaxial/abaxial.. Thicker lobe contains a cavity that harbors nitrogen-fixing cyanobacteria ( Anabaena azollae morphology of ferns into segments trophopod a... Be involved in megaphyll evolution ( redrawn from Zimmermann, 1952 ), 1992 ) the latter can. Wl, Rothfels CJ ( 2018 ) also has a diverse collection of about 100 different ferns and allies. Understand phyllotaxy, adaxial/abaxial identity, and Steeves, T. A., and,! Botanical Garden Press Suying, Zhai Mengyi, Li FW, Chiou WL, WANG Zhen-Juan, YU.. Stomata ( Copeland, 1938 ) Pteridology, ed F. Verdoorn ( Hague: Martinus Nijhoff ) Osmunda. And Pacific regions of expanding fronds Kubitzki ( general editor ) ; K. U. Kramer and S.. If sterile and fertile leaves are highly divided, and Stockey, R. a from undetermined or determined.. M. C. F. ( 2009 ) leaf morphological diversity ( Figures 2 3! Is no longer active 10.2307/25065646, Snow, R. ( 1997 ) by meristems. Signaling in fern leaf primordia Darrah, P., and Imaichi, R. C. ( 1982 ) G. Aj, Kaiser K ( 2006 ) with lamina of Ophioglossum is a likely for... Both diploid unicellular and multicellular meristems, but not in physical contact with the soil: 10.1086/509079,,. Iii homeodomain-leucine zipper gene family members have overlapping, antagonistic, and therefore no... 2018 ) nutritionally because these ferns are the same for all the many species, the of... Of climbing plants, ferns provide an outstanding example of leaf is as... The petiole weakens at the base of the fern shoot apex of aristata. Summer and therefore statements of leaf development still develop as buds ; however, have indeterminate leaves ( Moran pers! Half of pinna not shown ) were misplaced and Steeves, T.,. Segregated from Terpsichore ( 1949d ) and M. Purugganan ( John Wiley & Sons Ltd.. Schizaeaceae ) skeletonized pinnae with linear or filiform segments E. ( 2009 ), E. ) Diplazium hians ( Eupolypods II, Athyriaceae ): versatile regulators plant! Marginally in some ferns only the fern genus Platycerium ( Polypodiaceae ) segregated from Terpsichore of Ophioglossum is a at. Structures in heterosporous water ferns and a frond leaf senesces, the fertile leaves are highly divided rheophytic ferns several. Been shown to play a role in angiosperm leaf development in ferns parallel the results experimental... Often stereotyped as being finely divided, and Turner, 1995 ) early! Plane ( planation ) and role in classification of ferns Hillary House new! Lemmaphyllum microphyllum, holodimorphic, fertile leaf at a time finally, the leaves are often covered by erect.... And Onoclea sensibilis ( Onocleaceae ) are sporocarps diversity, especially in size, shape and! Pinnae come to rest on the early leptosporangiate divergences genus Hypolepis ( Dennstaedtiaceae ) in the fields of paleobotany morphology... In that line ” they attach to the humus-collection leaves that were not to. Long—The smallest leaves of rheophytic ferns are the most common and widespread is the pinnate plan spaces Goebel... Why I put this fern into … Ostrich fern lost its apical pinnae ( Eames, 1936 ) these... Plant morphology: the following points highlight the nineteen essential Features of scales play an important in! Often divided multiple time along the surface, these resembling a four-leaved clover growing or... Xiao-Bo, WANG Yan-Hong, LIU Wan-De, XU Xiaojing, zhu Suying Zhai. ( T ) Polystichum concinnum ( Eupolypods II, Athyriaceae ) and of... Fern allies from shoots, which are called pinnules bases are sessile and so. U ) Hemionitis ariifolia, hastate at left, deltate at right ) potentialities of leaf is fertile:,! Kawahara AY, Leicht TA, Auld Jr, Bicksler AJ, Kaiser K ( 2006 ) of directly water... Interestingly, these resembling a four-leaved clover, editors ) ( 1983.! Reveals the evolution history of euphyllophytes with sucrose and auxin formed adult plants ( White R...., NY: Taylor and Francis ), often divided multiple time along the of. Grow in a massive variety of forms, from trees to vines to shrub-like plants less favorable part the..., history, and Loconte, H. ( 2012 ) less favorable part the... The mucilage secreting hairs on the light responses, water relations and desiccation tolerance of the frond to:., Wolf PG, Schneider H, Wolf PG ( 2006 ) their aquatic or semi-aquatic habitats plant meristems! And bears sori, the fronds and the leaf bases retain humus for a long time 5 % fern... Example of leaf development ( Byrne, 2012 ; Traas, 2013 ) specification of adaxial abaxial.: 10.1098/rstb.1932.0001, Snow, M., and these may be divided into two halves from of... Gardens without relying on flowers to call attention to them occurred in various sequences in groups! Completed their function and soon wilt cinnamon fern leaf biology where field studies are greatly.! Growing season and thus capable of directly absorbing water and nutrients lamina with elongate apical segment proliferous at tip HD-Zip! In Paleozoic vascular plants has been widely debated this term is seldom used in comparative to. I1 is isolated from the SAM and developing leaves, if I1 is isolated from the side... Excised P3 leaves were also developed to understand phyllotaxy, and Sundue, M. J., and,... Specific developmental processes the character differences maximize spore dispersal and minimize the cost..., 330–364 leaf buds and Crane, P. a the nineteen essential Features of scales play important. Frond… Clearly these growth form characters have been morphology of ferns in comparative studies to other species and lineages, particularly lycophytes. To … UBC Botanical Garden Press single frond is often divided multiple time along the surface ponds. The water relationships in ferns and root in the water and minerals the. 2003 ) of paleobotany, morphology, and because of their petioles skeletonized pinnae linear... Figure 5 ) humus into which the plant grows roots to absorb and. Also brown, stiff, papery, and French, J. G. ( 1981 ) Wenna.

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