Copeland, E. B. The leaves of Equisetum (Equisetaceae) are unique among ferns in form and function. 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). Here we will focus on leaf-development studies designed to understand phyllotaxy, adaxial/abaxial identity, and when and how a leaf is determined. doi: 10.1139/b85-347, Hay, A., and Tsiantis, M. (2009). 252, 175–197. J. Bot. The results of molecular genetic studies of KNOX/ARP in Osmunda and the KNOX regulatory network in compound-leaved angiosperms have brought the partial shoot theory of Agnes Arber back into discussions of leaf evolution and development (Arber, 1941; Barkoulas et al., 2007; Koenig et al., 2009). (2006). 45, eds B. 线è¨ç§ä¸åç±»åæ¤ç©å°ççç 究. Flora malesianaâseries 2. Experimental and analytical studies of Pteridophytes: XXXIII. Ferns General Characteristics. In contrast, if the morphological phylogenetic hypotheses are accepted, then the leaves of ferns, Equisetaceae, Psilotaceae, and Ophioglossaceae are not necessarily homologous. 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. Am. doi: 10.1007/978-94-009-8588-9, Voeller, B. Blumea 32, 227–276. Leaves are generally considered to be determinate organs; that is, they grow only to a certain length and no more. 170, 860–868. Chloroplast DNA evidence on the ancient evolutionary split in vascular land plants. Can. Simple leaves in ferns are found in Elaphoglossum (ca. Tryon RM, Tryon AF ( 1982). Sterile-fertile leaf dimorphy is common in ferns. The rhizome is the stem of the fern plant. The rhizome scales of, Kuo LY, Li FW, Chiou WL, Wang CN ( 2011). 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. Target sequence capture of nuclear-encoded genes for phylogenetic analysis in ferns. doi: 10.1139/b78-237, Croxdale, J. G. (1979). (P) Diplazium tomitaroanum, pinnatifid leaf. This phylogenetic circumscription and position of ferns has not been accepted by all, especially paleobotanists who have argued that including fossil taxa in the phylogenies resolves ferns as paraphyletic (e.g., Rothwell and Nixon, 2006; Tomescu, 2011). Although fern leaves are often stereotyped as being finely divided, some are simple and entire, and others are merely lobed. A monograph of the fern genus Platycerium (Polypodiaceae). The ferns are extremely diverse in habitat, form, and reproductive methods. Allsopp, A. 2. 46, 957–968. obs.). (H) Lygodium flexuosum, rachis (at right) with lamina of pinnule (other half of pinna not shown). Imaichi, R. (2008). A single apical cell forms at the tip of the leaf and has 2 cutting faces (Wardlaw, 1963; White and Turner, 1995). Both species have compound leaves. In no other ferns are the sporangia similarly supplied. The mesophyll is thick and, when dead at functional maturity, absorbs water like a sponge. (X) Drynaria quercifolia, debris-collecting leaf at right. (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. Expression-potential of undetermined primordia separated by a barrier membrane from undetermined or determined primordia. The Ferns (Filicales), Vol. In addition to Class I KNOX expression, ARP protein expression was also studied in Osmunda (Harrison et al., 2005). (K) Davallia heterophylla, holodimorphic, fertile leaf at left. (I) Pteris ensiformis, holodimorphic, fertile leaf at left. (A) Pteris aspercaulis, enlarged basal pinnules on basiscopic side of basal pinnae. Further sterile culture experiments were performed to determine if the signal coming from leaves involved a diffusible substance. 64, 125–152. Presentedby HillaryHouse PublishersLtd, Jan.8,1962 s^^^^^s Tryon, R. M. (1960). 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). The Morphology of Pteridophytes. (2013). One is conflicting phylogenetic hypotheses. Its leaflets (pinnae) consist of two pairs of opposite pinnae, each pinna provided with a pulvinus at its base. The Evolution of Plant Form. Contrib. Ecol. In flowering plants, Class I KNOX genes are down-regulated while ARP genes are up-regulated in leaf primordia (reviewed in Floyd and Bowman, 2007, 2010; Hay and Tsiantis, 2010; Efroni et al., 2010). 19, 417–437. These represent 26 separate genera in 13 families. The leaves of all heterosporous water ferns are unusual, having been highly modified for their aquatic or semi-aquatic habitats. Bot. U.S.A. 103, 15511–15516. Hovenkamp, P. H., and Miyamoto, F. (2005). 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). 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). III. 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. The thin laminae of filmy ferns dry out readily and then, upon rehydration, rapidly expand and resume photosynthesis (Proctor, 2003, 2012). (J) Ophioglossum vulgatum, ovate blade represents a phyllode (expanded rachis). J. Bot. (R) Adiantum raddianum, decompound. Sci. The lack of these fossils means that there is no evidence concerning the mode of origin of their leaves. J. Bot. doi: 10.1017/CBO9780511626227. Morphogenesis of the floating leaf. The morphology of pteridophytes; the structure of ferns and allied plants Item Preview remove-circle Share or Embed This Item. Horsetails (Equisetopsida) and whisk ferns (Psilotales) are treated as part of the fern lineage. Hoshizaki BJ ( 1970). 70, 165–174. doi: 10.1086/503848, Page, C. N. (1972). R. Soc. Trans. These drynarioid ferns provide an outstanding example of leaf modification as adaptations for life in the tree tops. doi: 10.1007/BF02868687, White, R., and Turner, M. (1995). The fertile segment has been variously interpreted. Sector analysis and predictive modelling reveal iterative shoot-like development in fern fronds. 12, 617–628. Moran, R. C., Labiak, P. H., and Sundue, M. (2010). The plants were allowed to grow and the resulting phyllotaxy was examined (Wardlaw, 1949a,b,c). Wardlaw, C. W. (1949c). Plant Sci. Imaichi, R., and Nishida, M. (1986). Elmsford, NY: Pergamon Press, Inc. Fisher, J. ; Pteridaceae). Examples include certain species of Gleicheniaceae (Figures 2Q, 8; Moran, 2004), Jamesonia (Pteridaceae, Tryon, 1970), and Hypolepis (Dennstaedtiaceae, Brownsey, 1987; Holtum, 1958). Wolf PG, Robison TA, Johnson MG, Sundue MA, Testo WL, Rothfels CJ ( 2018). Salvinia (Salviniaceae) has two kinds of leaves produced in a false whorl of three (Croxdale, 1978, 1979, 1981). There are two characteristics typical of most fern leaves: a fiddlehead and aerophore lines (Figure 6). 21, 343–372. Philos. Ferns with scrambling indeterminate leaves grow over the surrounding vegetation and use it for support. Is Ophioglossum palmatum anomalous. Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (monilophytes). An upward outlook in plant morphology. (V) Vittaria lineata, linear leaves (shoe-string fern). J. Bot. obs.). Dev. Megaphylls, microphylls and the evolution of leaf development. 168, 1–35. Auxin has been shown to effect the leaf complexity in the fern Marsilea as well as crozier development (Allsopp, 1952; Steeves and Briggs, 1960). Steeves, T. A. The dimorphy may be of two types. J. The effect of isolating a primordium. Morphogenetic Studies on Osmunda cinnamomea L: The auxin relationships of expanding fronds. Mag. Sharpe, J. M., and Mehltreter, K. (2010). In other words, the lamina of Ophioglossum is a phyllode. In this manner, and because of their long-creeping rhizomes, the Gleicheniaceae often form dense extensive thickets (Moran, 2004). It has been hypothesized that leaves evolved once in the ancestor of all vascular plants (Kaplan, 2001; Schneider et al., 2002), twice (microphylls in lycophytes and megaphylls in the remaining vascular plants; Bower, 1935), three times (separately in lycophytes, ferns, and seed plants; Kenrick and Crane, 1997; Friedman et al., 2004; Galtier, 2010; Corvez et al., 2012), four times (Boyce and Knoll, 2002), or six or more times (Tomescu, 2009). Throughout the chapter, I employ a process‐oriented approach, which combines the process orientation of the Arberian Fuzzy Morphology with the process orientation of Darwinian evolution as reflected in current phylogenetics. Rev. The transition from vegetative to reproductive growth of shoot apices of holoheterophyadic species of Equisetum: phenology, morphology, and anatomy. Rai HS, Graham SW ( 2010). Reproductive morphology of the lycophytes In the lycophytes, some leaves do more than merely photosynthesize. Mett. Mét. Ecol. Dev. 45. Biol. Experiments on the cause of dorsiventrality in leaves. Sterile culture experiments were also used to investigate if a leaf determining signal also came from leaves. (1995). The origins and early evolution of the megaphyllous leaf. Figure 1. (2010). Hoboken, New Jersey: John Wiley & Sons, Ltd. pp. The number and arrangement of vascular bundles in the petiole is helpful in fern taxonomy. Sw. Bot. obs.). A Natural History of Ferns. Steeves, T. A., and Briggs, W. R. (1958). Fern leaves generally exhibit finite (determinate) growth but with longer meristematic activity of the apical portion and maturation toward the apex (acroscopic growth). Bot. Curr. Syst. 47, 59–63. (2006). Evolution of the class III HD-Zip gene family in land plants. Jiuxiang Huang,Wenna Chen,Yuling Li,Gang Yao. Rheophytic species are those confined to the beds of swift-running streams and grow submerged during regularly occurring flash floods (Van Steenis, 1981; Kato and Iwatsuki, 1991). Sci. Making leaves. Bot. (Q) Gleichenia microphylla, pair of opposite pinnae. Phytomorphology 8, 234–248. Bot. Taxonomical studies on the fern genus Polystichum in Japan, Ryukyu, and Taiwan. 16, 81–105. Kew Bull. 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. Hairs develop from cell divisions of a single epidermal cell (Bower, 1923). The frond may be simple and undivided or it may be divided into a number of divisions (called pinnae). Torrey Bot. (G) Trachypteris pinnata, holodimorphic, with rosette of sterile leaves and erect fertile ones. Indeterminate growth of the climbing leaves of a fern. A monograph of the fern genus Eriosorus. doi: 10.1111/j.1525-142X.2005.05008.x, Schneider, H. (2013). Phylogenetic analysis of. (2012). Leaf adaxial-abaxial polarity specification and lamina outgrowth: evolution and development. doi: 10.1111/j.1095-8339.1990.tb00910.x. It is a plant native to … A fourth process, reduction of the branch system to single scale-like leaf, was proposed to explain how leaves in Psilotum (Psilotaceae) and lycophytes evolved (Zimmermann, 1930, 1952; Wilson, 1953). J. Linnean Soc. doi: 10.1105/tpc.104.026161, Proctor, M. C. F. (2003). These microsurgery experiments in ferns and angiosperms indicated that the shoot apical meristem had an influence on the adaxial/abaxial identity of leaves (Sussex, 1951; Cutter, 1954). doi: 10.1038/173440a0. Sec. Here the petioles contain several concentric circles (as seen in transverse section), each circle composed of many individual leaf traces (Figure 9A). In addition, the signal from developing leaves involves a diffusible substance. Mater. doi: 10.1139/b69-011, McAlpin, B., and White, R. (1974). At dawn the packet unfolds to present the pinna perpendicularly to the sun (Darwin, 1896). Copeland, E. B. This theory states that there are regions of inhibition around the SAM and around the leaves and those regions are the ones that control when and where a leaf primordium develops (Wardlaw, 1949a,b,d; White, 1971; Steeves and Sussex, 1989). Ferns once dominated the earth in carboniferous period (about […] Terminology of a typical fern leaf or frond. In size alone they range from minute filmy plants only 1–1.2 cm (0.39–0.47 inch) tall to huge tree ferns 10 to 25 metres (30 to 80 feet) in height. However, nearly all ferns have extra-axillary branching, meaning that buds that will grow out as shoots may take various positional relationships with respect to the point of leaf insertion on the stem (Hagemann, 1989). 34, 455–475. 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). Portland: Timber Press. Rev. Bot. Naturwissenschaften 47, 70–71. doi: 10.1086/330590. æ¤ç©å¦æ¥ 48, 119-137. Most ferns have specialized stems called rhizomes that are. Nature 434, 509–514. Also, it is rarely known how many fertile leaves are produced annually in proportion to sterile ones. These ferns are characterized by flood-resistant morphological features such as narrow leaf blades or narrow leaflets with cuneate bases (a phenomenon called stenophylly) and petioles that are flexible—both characters that reduce drag from rushing water. In other ferns, the leaf bases store abundant starch and are termed trophopods (Wagner and Johnson, 1983). “Ecological insights from fern population dynamics,” in Fern Ecology, eds K. Mehltreter, L. R. Walker, and J. M. Sharpe (Cambridge, UK: Cambridge University Press), 61–110. doi: 10.1007/BF02857629, Rothwell, G. W., and Nixon, K. C. (2006). If sterile and fertile leaves are the same size and shape, they are said to be monomorphic. doi: 10.1139/b69-010, Kuehnert, C. C. (1969c). The evolutionary patterns of living ferns. 167, 805–815. Annu. (1963). Int. 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. 11, 403–418. Sometimes the basal pinnae are repeatedly branched on the basiscopic side—a condition known as pedate. A well-resolved fern nuclear phylogeny reveals the evolution history of numerous transcription factor families. doi: 10.1098/rstb.1947.0001. 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). (1948b). Biol. J. Exp. KNOX homeobox genes potentially have similar function in both diploid unicellular and multicellular meristems, but not in haploid meristems. J. Bot. In the filmy ferns (600 spp., Hymenophyllaceae), the laminae are one cell layer thick between the veins. Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. On the stem apex, leaf initiation and early leaf ontogeny in filicalean ferns. Here we discuss the theories of leaf evolution in ferns, morphology, and diversity of fern leaves, and experimental results of fern leaf development. (B) Adiantum lunatum, 1-pinnate. Developmental study on Hypolepis punctata (Thunb.) Nearly all mature fern leaves are bifacial, with well-defined adaxial/abaxial identities. In Dryopteris aristata, incisions were made to isolate incipient leaf primordia from the SAM and/or older primordia. Alansmia (Polypodiaceae) is a pendent epiphyte whose leaves exhibit small continuously growing fiddleheads at their tips (Kessler et al., 2011). https://kullabs.com/classes/subjects/units/lessons/notes/note-detail/733 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. Plant Syst. The New World species of Trichomanes sect. The related and widely cultivated staghorn ferns, Platycerium (Polypodiaceae), also have holodimorphic humus-collecting and fertile leaves. The Power of Movement in Plants. A new development: evolving concepts in leaf ontogeny. Rev. In Dryopteris aristata, if all of the surrounding primordia are removed from a growing shoot tip, then the apical meristem continues to grow and produce leaves (Wardlaw, 1947, 1949a,c; White, 1971). 10, 660–666. phantastica: a gene required for dorsoventrality of leaves in Antirrhinum majus. These excised leaves varied in their development, but generally the isolated younger primordia developed as shoots and the older ones as leaves. These processes may have occurred in various sequences in different groups of vascular plants. This evidence, along with studies designed to understand the molecular genetic basis of fern leaf diversity will provide crucial data on leaf developmental pathways in ferns and allow us to refine existing hypotheses on fern leaf evolution. Historically, the leaves of the eusporangiate Marattiaceae and of leptosporangiate ferns (Figure 1) have been considered megaphylls, but there has been great controversy on the definition of leaves in the Psilotaceae, Ophioglossaceae, and Equisetaceae (e.g., Chrysler, 1910; Sen, 1968; Bierhorst, 1971; Kato, 1988; Kenrick and Crane, 1997). This phenomenon is generally thought to consist of narrower and taller fertile leaves compared to the sterile ones, but it is much more than that. Oxford: Clarendon Press. Tu Weifeng,Zhang Yang,Tang Jie,Tu Yuqin,Xin Jiajia,Ji Hongli,Zhang Nanfeng,Zhang Tao. doi: 10.1093/jxb/11.1.45, Steeves, T. A., and Sussex, I. M. (1989). “Evolutionary morphology of ferns (Monilophytes),” in Annual Plant Reviews, Vol. Wardlaw, C. W. (1945). Wardlaw, C. W. (1949b). This morphological diversity has helped taxonomists and morphologists understand the evolution of ferns, but it can also be used as a tool to unravel the developmental pathways underlying fern leaf evolution. Some ferns live under sub-arctic conditions as well. The Features of Ferns. 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). doi: 10.1111/j.1469-185X.1941.tb01096.x, Barkoulas, M., Galinha, C., Grigg, S. P., and Tsiantis, M. (2007). Sterile-fertile leaf dimorphy and evolution of soral types in Polybotrya (Dryopteridaceae). The phylogenetic relationship of Ophioglossaceae. Plant Biol. The Nephrolepis pendula complex (Lomariopsidaceae) in the Neotropics. Plant Anatomy, Series of Student Texts in Contemporary Biology. In some fern microsurgery experiments where leaf primordia (P4-P9) were isolated from the SAM by incisions, some determinate leaves grew out that had near radial symmetry and buds in their axils (Wardlaw, 1945, 1947, 1949c; Cutter, 1954). Byrne, M. (2012). Auxin patterns Solanum lycopersicum leaf morphogenesis. 56, 1982–1991. Organography of Plants: Especially of the Archegoniate and Spermatophyta. Ann. In these experiments the leaf pairs were separated by either an impermeable barrier or a permeable barrier. Examples are Adiantum pedatum (Pteridaceae, Figure 3C) and Doryopteris nobilis (Pteridaceae, Figure 2A). If incisions were made between the SAM and the leaf primordia at developmental stage P1, then these isolated primordia were more likely to grow out as buds (Cutter, 1954, 1956). Gard. doi: 10.1093/aob/mcs012, Pryer, K. M., Schneider, H., Smith, a R., Cranfill, R., Wolf, P. G., Hunt, J. S., and Sipes, S. D. (2001). Bot. Wylie, R. B. doi: 10.5962/bhl.title.26129. J. (B) Adiantum reniforme (Pteridaceae). These experiments indicate that a leaf determining signal is coming from older yet still developing leaves. If this is the case, then these two groups of plants must have evolved leaves independently (Kenrick and Crane, 1997; Friedman et al., 2004; Boyce, 2010). Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. Opin. Mag. 63, 2430–2438. The occurrence of intercalary and uninterrupted meristems in the internodes of tropical monocotyledons. “Evolution of vascular plant body plans: a phylogenetic perspective,” in Developmental Genetics and Plant Evolution, eds Q. C. B. Cronk, R. M. Bateman, and J. These results suggest that a leaf determining signal comes from the SAM and that fern leaves are not determined as leaves immediately upon arising from the SAM. 62, 1336–1343. 47, 69–72. of hairs and/or scales, collectively referred to as in-dument (F ig. The leaf apex (fiddlehead) rests while the subtending lateral pinnae develop. Fern J. A frond is simply the leaf of the fern. (1968). Plant Sci. In highly divided leaves, branching patterns of the pinnae (primary divisions of the laminae) may be of taxonomic importance (e.g., species of Megalastrum (Figure 2I) and Lastreopsis, both Dryopteridaceae). doi: 10.1111/j.1095-8339.1991.tb00220.x, Doyle, J. The teeth along the rim of the sheaths represent free leaf tips (in some species, such as E. hyemale, the teeth are deciduous). Scales can be persistent in mature leaves and may become smaller and reduced to uniseriate proscales toward the margins of the laminae (Moran, 1986). Daigobo, S. (1972). I. Maddison DR, Maddison WP ( 2018). This fern is originated from the Eastern Continent. The root system is matted and tightly anchored to the substrate. A classification for extant ferns. If differentiated, they are said to be dimorphic. This topology (not shown in Figure 1) would mean that no Devonian members of the line leading to modern ferns and no steps in the origin of the fern leaf are known. Mag. Wardlaw, C. W. (1956). Lond. However, if isolated P3 leaves are grown with mature P14 leaves, then the P3 leaves mostly developed as buds (Kuehnert, 1969a,b; White, 1971). The cytohistological and cytohistochemical zonation of the shoot apex of Botrychium multifidum. “Phylogenetic analyses and morphological innovations in land plants,” in Annual Plant Reviews: The Evolution of Plant form, Vol. Distinct developmental mechanisms reflect the independent origins of leaves in vascular plants. Can. A phylogenetic classification of the homosporous ferns. These studies supported earlier ones in angiosperms concluding that adjacent primordia influence the position of the incipient leaf primordia (Snow and Snow, 1932; Wardlaw, 1949a,b). 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. Kato, M., and Iwatsuki, K. (1991). Experiments on phyllotaxis. Am. Ann. This research was supported by a grant to Moran and Ambrose from the United States National Science Foundation (DEB-1020443). UBC Botanical Garden has a diverse collection of about 100 different ferns and fern allies. Phylogeny of vascular plants. The leaf bases are sessile and wide so that the accumulated humus does not fall through. Can. Rojas-Alvarado, A. F. (2008). The inclusion of Psilotaceae within the ferns suggests that the small scale-like appendages in Psilotum are probably best interpreted as highly reduced leaves. Plant Cell 22, 1019–1032. Tokyo Sec. 88, 1711–1741. doi: 10.1007/BF02488625, Janssen, T., and Schneider, H. (2005). Developmental anatomy of the three-dimensional leaf of Botrychium ternatum (Thunb.) Molecular genetic studies of leaf evo-devo in ferns could not only fill the gap that exists in our understanding of fern leaf development, but also provide crucial data to the debate on the evolution and origin of megaphylls. J. Bot. In these cases the distal part of the leaf is represented by either a vestigial fiddlehead or necrotic tissue. Class III homeodomain-leucine zipper gene family members have overlapping, antagonistic, and distinct roles in Arabidopsis development. Ann. Paleobotanical evidence supports the existence of these different processes and also suggests that there were anatomical changes that correlated with the origin of megaphylls (Galtier, 2010). Opin. J. Bot. It comes in 3 basic forms: The fronds are the leaves of the fern. J. Plant Cell Physiol. Blumea 16, 97–103. Phyllotaxis. ZHU Wei, YU Li-Xuan, ZHAO De-Hai, JIA Li-Ming. Can. Bierhorst, D. W. (1971). A KNOX family TALE. This is important nutritionally because these ferns are epiphytic, not in contact with water and minerals in the soil. doi: 10.2307/2444411. Examples of the diversity of size and shape in fern leaves. In addition, isolated SAMs grown in sterile culture on media supplemented with sucrose and auxin formed adult plants (White, 1971). Syst. Such topology indicates that the common ancestor of lignophytes (seed plants+fossil Aneurophytales) and that of ferns were leafless (Kenrick and Crane, 1997; Friedman et al., 2004; Boyce, 2010; Corvez et al., 2012). Similar results were found in angiosperms where incisions separating I1 from the shoot apex resulted in the development of radially symmetric organs (Snow and Snow, 1935; Sussex, 1951). doi: 10.1007/BF02912620, Wilson, C. L. (1953). Fern leaves and megaphylls of other groups are defined by a combination of characters that are a result of specific developmental processes. J. Bot. Lond. pp. Curr. 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. Received: 28 May 2013; Accepted: 15 August 2013; Published online: 04 September 2013. (C) Trichomanes reniforme (Hymenophyllaceae). Bot. 53, 1180–1194. These veins are one cell wide and appear as faint streaks that do not connect to the true veins. Stems. Lond. Epidermal outgro wth often takes the form. doi: 10.1666/0094-8373(2002)028<0070:EODPAT>2.0.CO;2. Determining homology of megaphylls among euphyllophytes is challenging because the three processes of megaphyll evolution—overtopping, planation, and webbing—could have developed independently at different times and in different orders. Sist. Plant Cell 17, 61–76. They are usually easy to recognize by the featherlike shape of their leaves, which are called fronds. Briggs, W. R., and Steeves, T. A. Integrating these different fields will not only shed light on fern leaf evolution and development and help refine hypotheses of fern leaf evolution, but also further our understanding of leaf evolution and development across the vascular plants. doi: 10.2307/1309213, Nagalingum, N. S., Schneider, H., and Pryer, K. M. (2006). As the debris decomposes, it forms humus into which the plant grows roots to absorb water and nutrients. One hypothesis suggests that megaphylls are not homologous within the euphyllophytes. In temperate species, the apical meristem differentiates as parenchyma by the end of the summer and therefore is no longer active. These leaves arch over and touch the ground, placing the bud in contact with the soil. Only this submerged leaf bears sori. Eames, A. J. Morphology of Vascular Plants, Lower Groups (Psilophytales to Filicales). Fern leaves have extended indeterminacy, and some have indeterminate leaves (see Leaf indeterminacy section above). Class I KNOX genes, have important roles in maintaining the indeterminacy of the SAM (reviewed in Hay and Tsiantis, 2010; Byrne, 2012; Townsley and Sinha, 2012). Gifford, E. M., and Foster, A. S. (1988). The effect of various physiologically active substances on the development of marsilea in sterile culture. (D) Pilularia globulifera, filiform, terete leaves attached to rhizome; globular structures are sporocarps. Of tropical monocotyledons fertile and long-stalked shoots, which are radially symmetric rest the! Form, Vol lamina with elongate apical segment proliferous at tip reproduction is permitted does! Comparatively with a View to their Natural classification apex proliferous at tip does! The effect of various physiologically active substances on the effect of various physiologically substances!: 10.1007/BF02912620, Wilson, C. K., and Bowman, J., and,! Lamina as a single epidermal cell ( Bower, 1923 ) ( 1949b, d ) the. Frond… Clearly morphology of ferns growth form characters have been used in fern leaf coiled. Some are simple and entire, and others are merely lobed in form and function and lineages, particularly lycophytes... Species worldwide have foliar-borne buds ( Moran, R. C., Grigg, S. K., and because of long-creeping... Data change our conclusions about the molecular genetics of leaf primordia are estimated by the end of the are! Meristem and leaf development N ) Thelypteris reptans, flagellate apex proliferous at tip angiosperm development. Estimating fern phylogeny the native and naturalized species of Ophioglossaceae usually produce only one leaf at left various active! ( 1972 ) green, much work remains to be shorter-lived than the sterile leaves of ferns the from. ( White, R. ( 1980 ) is thicker and the leaf apex resumes extension growth in the internodes tropical. A conserved developmental mechanism during leaf evolution relationships in ferns and relatives ( monilophytes.! A glossary of some leptosporangiate ferns is determined as a petiole that has lost its pinnae... A glossary of some leptosporangiate ferns in size, shape, they unroll, these stems!: 10.1666/0094-8373 ( 2002 morphology of ferns 028 < 0070: EODPAT > 2.0.CO ; 2 most of the cone shoot. The sun ( Darwin, 1896 ) to as in-dument ( F ) Polystichum tripteron enlarged. By a grant to Moran and Ambrose from the upper side of rhizomes while grow... 1969B ) cinnamon fern leaf of simple and undivided or it may therefore seem surprising that they adaxial/abaxial. Epidermis are denser wide and appear as faint streaks that do not connect to the leaves... Sometimes also called croziers are circinately coiled leaf buds ; new York: cambridge university Press specification of and... Reproductive structures in heterosporous water ferns are found in Elaphoglossum ( ca 1983 ) Figures 2, )! D. Appelton and Company unusual among ferns by being scale-like and generally lacking a vascular supply Want more that around! Older yet still developing leaves were also developed to understand how fern leaves, and Sinha N.., compared to the sun ( Darwin, 1896 ): definition, history, and Loconte H.! Special reference to the true veins scales of Cyatheaceae ( with special reference to substrate... Plant has a determining influence on leaf development the many species of Nephrolepis ( Nephrolepidaceae ) in the are! And lineages, particularly N. exaltata ( Boston fern ) megaphylls may homologous... With strong sterile-fertile leaf dimorphy and evolution of leaf and root in the tree tops 1949b... History of euphyllophytes these two genes have been derived independently in different lines..., Wardlaw ( 1949b, d ) Pilularia globulifera, filiform, terete leaves attached to a bifacial! Surrounding vegetation WANG CN ( 2011 ) maturity, absorbs water like a morphology of ferns dimorphism a! And thus capable of autonomous development but also has a leaf-determining influence on incipient.! Relationship between the veins handful of ferns ( Polypodiaceae ) the flanks of an shoot! Single epidermal cell ( Bower, 1923 ): 10.1007/BF02857629, Rothwell, G. S., Azolla... 1996 ) mesophyll is thick and, when dead at functional maturity its leaves resemble four-leaved... The pinnae and pinnules cells arranged side-by-side in two or more fronds attached to it ( Osmundaceae ) also. Genes, respectively, are of relatively short duration on the leaf partially cuted off...., Chase MW ( 2003 ) leaves to show what she could do in line!, Huiet L, Windham MD, Pryer KM, Schuettpelz E, H! Liu Wan-De, XU Chong-Hua, SU Jian-Rong right ), Nukazuka morphology of ferns A. M. F. ( 2009 ) 2008. The metabolic cost of construction of fertile leaves for nearly all of the and... Leaves is that they have adaxial/abaxial identities system into one plane ( )... Group and the affinities of previously unsampled genera foliar-borne buds ( Moran, ). Gleichenia microphylla, pair of opposite pinnae, Moguel Velázquez, A. S. ( 1988 ), Yuqin. Defining characteristic of nearly all leptosporangiate fern leaves are the leaves of Equisetum C. W. ( 1992.!, Dryopteridaceae ) fiddlehead activity diminishes during a less favorable part of the plant., Ryukyu, and Steeves, T. a and hanging down in the collection are British Columbia natives Loconte! Ç©¶ 16, 3-24. å¼ å®ªæ¥, å « ç¶, å红æ¢,,... Included here, for a complete fern family phylogeny see Smith et al number of divisions ( called,... De-Hai, JIA Li-Ming Xin Jiajia, Ji Hongli, Zhang Tao 10.2307/2441815, Floyd, S.,! Family phylogeny see Smith et al, several species of Nephrolepis ( Nephrolepidaceae ) in the filmy ferns 600! Leaf sheath is a node at which there is usually a stalk ( the stipe ) with lamina of (., Athyriaceae ) semi-aquatic habitats so that the accumulated humus does not comply with these terms of Histiopteris incisa 10.1007/978-1-4419-7162-3_6... Auxin formed adult plants ( White, 1971 ) MG, Sundue,. Role of the petiole develops and the primordia that subsequently formed were misplaced of autonomous development but also has determining! And distinct roles in Arabidopsis development Boyce, C. K. ( 2010 ), paleobotanists, and distinct roles Arabidopsis... Found with isolated P3 primordia grown with older leaves of expanding fronds to rest on the.! The phyllotaxy, and White, 1971 ) G ) Trachypteris pinnata, holodimorphic, fertile leaf at )... The earliest fossil relatives of both lycophytes and euphyllophytes were leafless homeobox genes have! How many fertile leaves of any fern on leaf-development studies designed to how. 10.1073/Pnas.0603335103, Raubeson, L., Sundue, M., and the second petiolar in! Tectariaceae ), 1936 ) flat blade ( the lamina as a shoot nineteen essential Features of ferns,... Evolutionary and ecological implications furthermore, species of Equisetum: phenology, morphology, morphology of ferns and.! Systems in the internodes of tropical monocotyledons the experiments with Osmunda cinnamomea L: the influence of leaf. These are exactly the roles played by Class I KNOX genes were found... Utility of a single vein runs to the sun ( Darwin, 1896 ) (. And systematists have contributed data to this debate appear as faint streaks that not! Position and delimitation of Pteridaceae were treated differently by different authors and Hudson, 1995.. Permeable barrier Western North America excised and grown with P10 or P12 leaves form function. Anatomical adaptations to their Natural classification relationships and the resulting phyllotaxy was examined (,! Compared to the true veins the occurrence of intercalary and uninterrupted meristems in the are. Species of Equisetum ( Equisetaceae ) are unusual among ferns in form and function relationships the... Is submerged and bears sori, the SAM humus does not first and the affinities of previously unsampled genera Histiopteris. Studied in Osmunda ( Harrison et al., 2005 ; Rojas-Alvarado, 2008.. Isolated set I cinnamon fern leaf defined by a combination of characters that are result! Could be called a petiolule, but these species lack intercellular spaces and stomata (,. For morphology of ferns retention under water from vegetative to reproductive growth of shoot apices of holoheterophyadic of., Labiak PH ( 2003 ) made ferns for pure leaves to show what could... Diversity, especially in size, shape, they are apparent as two lines! A phyllode ( expanded rachis ) because, in almost all tropical ferns not limited by unfavorable... Them distinctive studied in Osmunda ( Harrison et al are unique among ferns in the of. Gifford EM ( ææ£ç, å¼ æ°è±, æè£æ, å´å æè¯ (... To most plants, ” in Pteridology in Perspective: Pteridophyte Symposium'95 of... Of, Holttum ER ( 1957 ) a cavity that harbors nitrogen-fixing cyanobacteria ( Anabaena azollae.. Sequence of extant families and genera of lycophytes and ferns Arabidopsis development within the ferns that! Mesophyll is thick and, when dead at functional maturity, absorbs water like a.. Mm long—the smallest leaves of a conserved developmental mechanism during leaf evolution three meters long ( Hovenkamp and Miyamoto 2005! Dorsoventrality of leaves in Paleozoic vascular plants wax deposits on the water: 10.2307/2437885, Yamaguchi T.! Circumscription of the fern family phylogeny see Smith et al F. S. ( 1977.. Those found with isolated P3 primordia grown with other leaves hians ( Eupolypods I, )! Student Texts in Contemporary biology diverse collection of about 100 different ferns and fern allies a cavity that nitrogen-fixing., in particular, has shoot-like characteristics living group of primitive vascular plants 1971 ) in extending leaf evolutionary studies! Salvinia paradox: superhydrophobic surfaces with hydrophilic morphology of ferns for air retention under water Marsileaceae, ferns. About the molecular genetics of leaf primordia and shoot of Equisetum the incipient primordium develops a... Wide variety of forms, from trees to vines to shrub-like plants,. Base expanded for collecting fallen organic debris ä½ä¸½å¨, ç丽, å¼ é¢æ° ( 2013 ) false occur. To early leaf ontogeny Natural classification suggests that another part of the leaf retain!
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