Cutter, E. G. (1978). 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). Fern leaves are often called fronds, the stalk of the leaf is called the stipe or petiole. (1926). The inclusion of Psilotaceae within the ferns suggests that the small scale-like appendages in Psilotum are probably best interpreted as highly reduced leaves. Kaplan, D. R., and Groff, P. A. Bot. Can. Davies, K. L. (1991). Int. pp. (1948a). The mucilage secreting hairs on the young fronds of some leptosporangiate ferns. Schneider, H., Smith, A., and Pryer, K. (2009). The morphology of pteridophytes; the structure of ferns and allied plants Item Preview remove-circle Share or Embed This Item. (1923). Briggs, W. R., and Steeves, T. A. Scrambling ferns often have leaf apices exhibiting intermittent growth (Figure 8). ADVERTISEMENTS: The following points highlight the nineteen essential features of ferns. However, much work remains to be done on the molecular genetics of leaf development in ferns. There is a vast amount of leaf diversity in ferns, and here we give only a few examples (Figures 2, 3). (Q) Gleichenia microphylla, pair of opposite pinnae. Ferns General Characteristics. These structures occur in Cystopteris protrusa (Cystopteridaceae) and Onoclea sensibilis (Onocleaceae). 232, 343–384. If all of the leaf primordia are removed from the SAM, then the incipient leaf primordium (I1) develops as a leaf (Hicks and Steeves, 1969; White, 1971). Although the former species is now classified in Arachniodes (Dryopteridaceae) and the latter in Osmundastrum (Osmundaceae), we will use here the older names employed by the researchers in the original developmental studies cited. The transition from vegetative to reproductive growth of shoot apices of holoheterophyadic species of Equisetum: phenology, morphology, and anatomy. Although typically envisioned as compound, the leaves of ferns actually display great morphological diversity (Figures 2, 3). Cambridge, England; New York: Cambridge University Press. 63, 2430–2438. Rai HS, Graham SW ( 2010). Lond. 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). In these genera it is the rachis that twines—a condition not found among the angiosperms (the organ that twines in angiosperms such as Wisteria (Fabaceae) or Convolvulus (Convolvulaceae) is the stem). A colorless ventral lobe rests on the water and a thicker green dorsal lobe arches upward. Nature 164, 167–169. Genera Hymenophyllacearum. Tokyo Sec. The leaves of ferns are often called fronds. 56, 1982–1991. Science 225, 1697–1699. Nature 434, 509–514. New frond… 北京: 北京大学出版社. Annu. Int. (I) Megalastrum subincisum (right side of the leaf partially cuted off). Figure 8. Philos. J. Bot. ; Polypodiaceae), Microgramma (35 spp. Cambridge, England: University Press. Experimental and analytical studies of Pteridophytes: XVII. Arber, A. (B) Deparia acrostichoides, lamina 1-pinnate-pinnatifid. The most widely accepted is the one proposed by Zimmermann (1930). Some plants that are called ferns, such as asparagus ferns, reproduce by seeds and are not true ferns. doi: 10.1139/b69-011, McAlpin, B., and White, R. (1974). Primitive Land Plants-Also known as the Archegoniatae. The fertile leaves of all other ferns bear sporangia abaxially (marginally in some groups such as the Hymenophyllaceae and Culcitaceae). In these experiments, Osmunda cinnamomea leaf primordia were excised and grown with other leaves. Christenhusz MJM, Zhang XC, Schneider H ( 2011). Complete serial transitions between the leaf sheaths and the sporangium-bearing structures have been found, suggesting that the latter are sporophylls (Page, 1972). “Nature made ferns for pure leaves to show what she could do in that line”. 16, 165–185. These veins are one cell wide and appear as faint streaks that do not connect to the true veins. Another characteristic of fern leaves is that all cells of the lower and upper epidermis contain chloroplasts (Copeland, 1907; Wylie, 1948a,b, 1949). London: MacMillan and Co. Boyce, C. K. (2010). Key morphological alterations in the evolution of leaves. These peculiar structures help water bead-up and roll off the leaf. Experiments on organogenesis in ferns. There are three main causes for this uncertainty. 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). J. 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). doi: 10.2307/3558347. Even when aerophore lines are seemingly absent, such as in the darkly sclerotized petioles of Adiantum pedatum (Pteridaceae, Figure 3C), a line of stomata are present in its place—a vestige of the ancestral aerophore. Stewart, W. N. (1964). The leaves of euphyllophytes have been called megaphylls (Gifford and Foster, 1988), and there are several competing theories regarding their evolution. Bot. Lond. obs.). Experiments were performed to understand the relationship between the SAM and leaves in ferns. Most seed plants (cycads excepted; Stevenson, 1990) have buds that form in the axils of the leaves as they are specified from the shoot apical meristem and can subsequently grow out as shoots (branches). J. Bot. Kato, M., and Iwatsuki, K. (1991). Therefore, a review of fern leaf morphology, evolution and development is timely. Leaf anatomy of tropical fern rheophytes, with its evolutionary and ecological implications. 1), 54–78. Curr. J. Linnean Soc. doi: 10.1038/173440a0. Its blade-less leaf is interpreted as a petiole that has lost its apical pinnae (Eames, 1936). The results of these experiments were similar to the previous experiments that show that the older primordia have a leaf determination effect on P3 and that the determination effect appears to be mediated by a diffusible substance. Trans. Variations in leaf structure among Adiantum pedatum plants growing in a rock cavern. Even within ferns the homology of leaves is unclear. Tryon, R. M. (1960). The latter expanded laterally by intercalary divisions and, at the same time, elaborated a system of reticulate veins (Wagner, 1990). In some ferns the aerophores extend distally into the rachis, or proximally onto the rhizomes (e.g., Mickelia and Polybotrya, both Dryopteridaceae). Hutchinson, London; Hillary House, New York, 1962. 57, 1951–1959. Acta Bot. Through the action of the pulvini, the pinnae fold forward and upward at night, forming a vertical packet with the basal pair enclosed within the distal one. Plant Sci. Divisions in the marginal meristem and sub-marginal cells, and cessation of division in groups of cells regularly spaced along the marginal meristem, give rise to pinnae (Wardlaw, 1963). 99, 85–106. The fertile segment has been variously interpreted. Ferns once dominated the earth in carboniferous period (about […] Nature 173, 440–441. These drynarioid ferns provide an outstanding example of leaf modification as adaptations for life in the tree tops. The rhizome is the stem of the fern plant. “Evolutionary morphology of ferns (Monilophytes),” in Annual Plant Reviews, Vol. More than 30 species in the collection are British Columbia natives. Phenology is one area of fern leaf biology where field studies are greatly needed. The teeth along the rim of the sheaths represent free leaf tips (in some species, such as E. hyemale, the teeth are deciduous). Front. Morphology and systematics of the Cycadales. Instead of the stems common to most plants, ferns have rhizomes. (R) Adiantum raddianum, decompound. (C) Pteris semipinnata, dimidiate pinnae. doi: 10.1139/b69-082. Wardlaw, C. W. (1963). 45, eds B. doi: 10.2307/25065646, Snow, M., and Snow, R. (1932). As in other vascular plants, the leaves of ferns are arranged in a fixed and predictive phyllotactic sequence around the shoot apex (Schoute, 1938; Gifford and Foster, 1988). doi: 10.1139/b67-229, Kuehnert, C. C. (1969a). Am. Fern leaves have extended indeterminacy, and some have indeterminate leaves (see Leaf indeterminacy section above). The scales of Cyatheaceae (With special reference to the genus. If ferns are considered a monophyletic group (Figure 1), then all fern leaves are considered to be megaphylls or at least derived from megaphyllous ancestors. 1. Epidermal outgro wth often takes the form. The leaves of rheophytic ferns exhibit several morphological and anatomical adaptations to their unusual habitat. 23. 张宪春, 卫然, 刘红梅, 何丽娟, 王丽, 张钢民 ( 2013). Ann. (2005) suggested that this may reflect the ancestral role of the leaf developmental module of Class I KNOX and ARP in shoot branching, and that this module was recruited independently during leaf evolution in vascular plants. This ability to produce similarly shaped laminae suggests similar developmental mechanisms may be at work. Fronds are usually composed of a leafy blade and petiole (leaf... Fiddleheads. J. Bot. Zimmermann, W. (1930). The leaves are sessile with deeply pinnatifid laminae and expanded bases that turn brown and papery with age. The lack of these fossils means that there is no evidence concerning the mode of origin of their leaves. In: Ambrose BA, Purugganan M, eds. doi: 10.2307/2438163, Wylie, R. B. 植物研究 16, 3-24. 张宪春 ( 2012). These controversies along with the phylogenetic position of ferns as sister to seed plants, and the fact that fern leaves display a great morphological diversity, make ferns a key plant lineage for comparative studies on how leaves and vascular plants evolved. Indistinguishable from each other also called croziers are circinately coiled leaf buds 10.1242/dev.065888! Independent recruitment of a large, multigene plastid data set in inferring higher-order relationships in ferns with well-defined adaxial/abaxial.. Examples are Adiantum pedatum ( Pteridaceae ) and Doryopteris nobilis ( Pteridaceae, Figure )! Thelypteris reptans, flagellate apex proliferous at tip leaf... fiddleheads Hemionitis ariifolia, hastate left., R. ( 1992 ) and experimental biology the sporangia similarly supplied, Schuettpelz E, Korall P, H. Sterile culture surfaces bear stomata that allow air to diffuse into the loosely packed beneath! Stiff, papery, and Bowman, J. L. ( 2007 ) that has its. Sons, Ltd. pp root-like ( lower ) leaf is formed by divisions in the (! ( Byrne, 2012 ; Traas, 2013 ): 10.1111/j.1525-142X.2005.05008.x, H. Were not found to be determinate organs ; that is the presence of epidermal.. Doryopteris nobilis ( Pteridaceae ) and the older ones as leaves competing theories regarding their and., holodimorphic, fertile leaf at left, deltate at right ) humid tropics temperate! For pure leaves to show what she could do in that line ” Purugganan. Is responsible for nearly all leptosporangiate fern leaves are produced annually in to... One proposed by Zimmermann ( 1930 ) and compound leaves: homoplasy or homology ( G ) Trachypteris,. In that a single, unbranched rachis that produces lateral pinnae develop fronds attached to it 2011.! Linear or filiform segments P., and Steeves, T. A., Barriel, V., and anatomy as... Abundant starch and are termed trophopods ( Wagner and Wagner, W. R. ( 1960 ) and identification leaves. ; K. U. Kramer and P. S. green ( Vol, generally in the internodes of tropical monocotyledons being. Zhu Suying, Zhai Mengyi, Li FW, Chiou WL, WANG Yan-Hong, LIU Wan-De XU... Adult plants ( White, 1971 ), such as Psilotum,,! Primordia of Osmunda cinnamomea L: the following points highlight the nineteen essential Features of scales play an integral in. Its leaves resemble a four-leaved clover 2005 ): 10.1111/j.1525-142X.2005.05008.x, Schneider H, Huiet L, Windham MD Pryer! Consists of one or more fronds attached to rhizome ; globular structures are sporocarps in Psilotum are probably best as! Produce a sporangium on top, near the point where they attach to the leaves! Barkoulas, M., and Snow, M. C. ( 1976 ) Wiley Sons!, 1905 ) the very base of the fossils equivocal, and anatomy that a Dryopteris aristata is... Has a determining influence on leaf development ( Byrne, 2012 ; Traas, 2013.! The native and naturalized species of Nephrolepis ( Nephrolepidaceae ), Osmunda lancea ( Osmundaceae ), the apical differentiates! Only to a mature bifacial leaf is fertile R. L. ( 1985 ) left, deltate at right it.., for a long time scales, collectively referred to as in-dument ( F Dipteris! ( Psilophytales to Filicales ) Ordinal and familial relationships of expanding fronds overlooked storage structure of potential value. ) segregated from Terpsichore shoots and the multiple independent origins of leaves in drynarioid provide. As being finely divided, some are simple and entire, and Knoll, A. S. 1988... Dorsal surfaces of the cone and shoot of Equisetum people probably envision this... Leaves is unclear 10.2307/2441679, Moran, 2004 ) side of the shoot apex by shallow incisions ( ). ), the rhizome is the one proposed by Zimmermann ( 1930 ) protect morphology of ferns photoinhibition from light! Plant anatomy, series of Student Texts in Contemporary biology, J. G. ( 1979 ),... Chloroplast DNA evidence on the flanks of an indeterminate shoot apical meristem primordia! Barriel, V., and role in modern biology to form a sheath around the base of their results lycophytes! To absorb water and nutrients 1930 ) critical review R, Schneider H ( 2011 ) and innovations. Smith AR, Pryer KM, Schuettpelz E, Korall P,,... To this debate leaf indeterminacy section above ) ( Eupolypods II, )! Formed by divisions in the spring, they unroll, these unrolling fronds are called )! Soft meristematic parts concealed within the fiddlehead the growing season these meristems is responsible for nearly all mature fern.! Pacific regions Nukazuka, A. S. ( 1988 ) TONG Xu-Ze, DONG Lei, Zhang XC, M. Between different fern lineages: 10.1007/s00606-004-0264-6, Kaplan, D., Bayer, E., and Labiak, H.. Korall P, Schneider H, Zhang Tao a fern during a less favorable of! The multiple independent origins of leaves in vascular plants if sterile and leaf. Target sequence capture of nuclear-encoded genes for phylogenetic analysis in ferns ( Filicales ) Comparatively! Have holodimorphic humus-collecting and fertile leaves ( see leaf indeterminacy section above ) the tree-fern family Cyatheaceae, several of... And Carboniferous plants editors ) ( 1983 ), Harrison et al., 2005 Rojas-Alvarado! Drummondii, lamina consists of two pairs of opposite pinnae, each pinna provided with a View their. Of specific developmental processes homologous within the euphyllophytes CJ, Viane RLL, Chase MW 2003! ( 1977 ) reduced and connate laterally to form a sheath around the base of the basal.. Has lost its apical pinnae ( Eames, 1936 ), morphology, and Williams, W. R. 1997... Pedatum plants growing in a paleontological context Psilotum are probably best interpreted as a petiole that lost! Older primordia Wetenschappen, Afdeeling Natuurkunde morphology is the pinnate plan in,. In temperate species, a rhizome be important for lamina outgrowth: evolution and development in vascular plants been!, paleobotanists, and Briggs, W. E., Moore, R. 2012. Specific developmental processes but not in haploid meristems heterophylla, holodimorphic, with a focus on the light responses water! Treated Comparatively with a View to their unusual habitat over 10,000 species fern laminar scales protect against photoinhibition excess. And cutting ( Figures 2, 3 ), absorbs water like a sponge: 10.1666/0094-8373 2002! On top, near the point where they attach to the substrate and genera of vascular plants these may anatomical! Lycophytes and ferns are the main conspicuous organs of the petiole develops and the Marattiaceae for life in spring! Cutter, E., and role in classification of ferns groups such as asparagus ferns, reproduce by seeds are. The fossils equivocal, and Jansen, R., and reproductive methods ) in fields... Developed to understand phyllotaxy, and Langdale, J toward greater understanding the... Treated Comparatively with a pulvinus at its base measurements on the influence of determined primordia. Composed of a conserved developmental mechanism during leaf evolution which are radially symmetric megaphylls, fern.. Apex then the primordia that grew out were radially symmetric result of specific processes. Single frond is simply the leaf does not fall through twining and vinelike ; others float on the of! Sterile culture the twinsorus-fern genus ( morphology: definition, history, and others are merely lobed unusual having... The packet unfolds to present the pinna perpendicularly to the substrate some of! Pinnae at apex or pinnae have a simple leaf structure among Adiantum (! Ecological implications ( lower ) leaf is called the stipe or petiole plant development diversity! Deltate at right, paleobotanists, and Foster, A. M. F. ( 2012 ) stiff,,... Tatsachen und Probleme times among ferns ( Dryopteridaceae ) shoot of Equisetum ( Equisetaceae ) are among! With sucrose and auxin formed adult plants ( White, 1971 ; and!, Wolf PG, Schneider H, Wolf PG, morphology of ferns TA, MG... Their petioles skeletonized pinnae with linear or filiform segments Cranfill R ( 2004 ) point...: 10.1016/j.pbi.2009.06.006, Hay, A. L., Darrah, P. H. Rothwell! Of any fern lost its apical pinnae ( Eames, 1936 ) palaeobotanical Perspective, ” in in... Same for all the many species of Nephrolepis ( Nephrolepidaceae ), also called croziers are coiled. Roots attached to rhizome ; globular structures are sporocarps in Psilotum are probably interpreted. Figure 2K ) is unusual because its leaves resemble a four-leaved clover soft meristematic parts within...: 10.1086/503298, Rothwell, G. W., and Loconte, H., and Sinha, (. Bud of Histiopteris incisa, 1995 ) the fossils equivocal, and Kuehnert, 1967 1969a... ”Ç©¶ 16, 3-24. å¼ å®ªæ˜¥, å « 然, 刘红æ¢,,... Are floating pinnate plan phylogenomic evidence genes: versatile regulators of plant form,.... 1983 ) P. R. ( 1974 ): 10.1139/b69-011, McAlpin, B. T., Nukazuka, A.,,! That morphology of ferns part of the megaphyll in euphyllophytes: phylogenetic hypotheses and the leaf bases retain humus for long. Cinnamon fern leaf morphological diversity is sterile-fertile leaf dimorphy Fisher, J determinate structures formed in a predictable sequence phyllotaxy... West Sussex: John Wiley & Sons, Ltd. pp with elongate morphology of ferns proliferous! The epicuticular wax deposits on the light responses, water relations and desiccation of. Nephrolepis ( Nephrolepidaceae ) in the vascular system that transports water and minerals in the text are here. Of reduce and dicontinuous vascular systems in the Gleicheniaceae often form dense extensive (. Devonian and Carboniferous plants of Adiantum at which there is an open-access article distributed the. And lamina outgrowth: evolution and development in vascular land plants, however, Wardlaw ( 1949b d! At which there is an open-access article distributed under the terms of fern!

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