Is morphology really at odds with molecules in estimating fern phylogeny. Most commonly, as the leaf senesces, the petiole weakens at the base, and the leaf gradually reclines toward the ground. Further studies on the influence of determined leaf primordia on undetermined leaf primordia. Morphogenesis of the floating leaf. Examples are Deparia acrostichodes (Thelypteridaceae, Figure 2B), Matteuccia struthiopteris (Onocleaceae, Figure 2M), and Megalastrum subincisum (Dryopteridaceae, Figure 2I). |, The Morphological Diversity of Fern Leaves, Experimental Analyses of Fern Leaf Development, Molecular Genetics of Fern Leaf Development, Creative Commons Attribution License (CC BY), The New York Botanical Garden, Bronx, NY, USA. The scientific name of ostrich fern is Matteuccia struthiopteris. The experimental induction of buds from leaf primordia in Dryopteris aristata Druce. Figure 5. Plant Cell 22, 1019–1032. 24, 1–18. Plant Biol. doi: 10.1016/j.pbi.2009.10.001, Boyce, C. K., and Knoll, A. H. (2002). doi: 10.2307/2441679, Moran, R. C. (1986). 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. Studies of the effects of homogenized, determined leaf primordia on expression-potential of undetermined leaf primordia. Schneider, H., Smith, A., and Pryer, K. (2009). These peculiar structures help water bead-up and roll off the leaf. Morphology and Anatomy. There is usually a stalk (the stipe) with a flat blade (the lamina), often divided into segments. (1995). 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). doi: 10.1086/503848, Page, C. N. (1972). This is important nutritionally because these ferns are epiphytic, not in contact with water and minerals in the soil. Fern leaves have extended indeterminacy, and some have indeterminate leaves (see Leaf indeterminacy section above). 63, 535–562. J. Bot. UBC Botanical Garden has a diverse collection of about 100 different ferns and fern allies. doi: 10.1002/9781118305881.ch1. Developmental potentialities of leaf primordia of Osmunda cinnamomea. PPG I ( 2016). The stalk of a pinna could be called a petiolule, but this term is seldom used in fern taxonomy. Rev. Waites, R., and Hudson, A. A single apical cell forms at the tip of the leaf and has 2 cutting faces (Wardlaw, 1963; White and Turner, 1995). A colorless ventral lobe rests on the water and a thicker green dorsal lobe arches upward. Its blade-less leaf is interpreted as a petiole that has lost its apical pinnae (Eames, 1936). Ann. In addition, if I1 is isolated from the apex then the incipient primordium develops as a shoot. Cutter, E. G. (1978). Pryer KM, Schuettpelz E, Wolf PG, Schneider H, Smith AR, Cranfill R ( 2004). Harvard Univ. 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. Fern laminar scales protect against photoinhibition from excess light. Key morphological alterations in the evolution of leaves. These pinnae, called aphlebiae, are of unknown function. A rhizome is a specialized, root-like stem. Morphogenetic Studies on Osmunda cinnamomea L: The auxin relationships of expanding fronds. 23. å¼ å®ªæ¥, å«ç¶, åçº¢æ¢ , ä½ä¸½å¨, çä¸½, å¼ é¢æ° ( 2013). The field of developmental genetics is the missing piece and when integrated with data from other fields will help us develop more robust hypotheses of fern leaf evolution and development and more broadly hypotheses of leaf evolution and development in vascular plants. The science of plant morphology: definition, history, and role in modern biology. Figure 2. Experimental induction of buds from fern leaf primordia. 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. The effect of isolating a primordium. These lines aerate the leaf. 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. Development 138, 2925–2934. (I) Megalastrum subincisum (right side of the leaf partially cuted off). 63, 2430–2438. ; Pteridaceae). More than 30 species in the collection are British Columbia natives. Am. Tsutsumi C, Kato M ( 2008). These results are similar to those found with isolated P3 primordia grown with P10 or P12 leaves. In nearly all extant ferns, leaves constitute the dominant organs of the plant. 3(Suppl. doi: 10.1126/science.1070343. 13, 102–107. A special case of fern leaf morphological diversity is sterile-fertile leaf dimorphy. (L) Olfersia cervina, holodimorphic, fertile leaf at right. (G) Trachypteris pinnata, holodimorphic, with rosette of sterile leaves and erect fertile ones. Distinct developmental mechanisms reflect the independent origins of leaves in vascular plants. 115-140. (1981). doi: 10.1007/978-1-4419-7162-3_6, Townsley, B. T., and Sinha, N. R. (2012). Most ferns are leptosporangiate ferns… 600 spp. Hutchinson, London; Hillary House, New York, 1962. Both species have compound leaves. In general, leaves are the main conspicuous organs of vascular plants and often easy to recognize as such. doi: 10.1038/167651a0, Tomescu, A. M. F. (2009). (1884). *Correspondence: Barbara A. Ambrose, The New York Botanical Garden, 2900 Southern Blvd., Bronx, NY 10458-5126, USA e-mail: email@example.com, Front. However, outside of the seed plants, comparative molecular studies so far have focused on only two transcription factors: Class I KNOX and Class III HD-Zip (Harrison et al., 2005; Prigge and Clark, 2006; Floyd and Bowman, 2006). One extreme is the polycyclic condition found in Acrostichum (Pteridaceae) and the Marattiaceae. doi: 10.1098/rstb.1949.0001, Wardlaw, C. W. (1949d). 55, 223–231. 200, 54–174. Variations in leaf structure among Adiantum pedatum plants growing in a rock cavern. doi: 10.1242/dev.065888, Sanders, H., Rothwell, G. W., and Wyatt, S. E. (2009). The root system is matted and tightly anchored to the substrate. doi: 10.1126/science.255.5052.1697. Genera Hymenophyllacearum. Can. Leaf Divisions. The dominant role of the epidermis in leaves of Adiantum. B Biol. (1995). Fern leaves arise from the periphery of a shoot apical meristem (SAM) in a distinct phyllotaxy as do seed plant leaves (Wardlaw, 1963; Gifford and Foster, 1988). A. Philipp. Schneider, H., Pryer, K. M., Cranfil, R., Smith, A. R., and Wolf, P. G. (2002). Only this submerged leaf bears sori. 65, 188–218. The surface of the leaf, except for those apical cells of hairs united by their tips, bear a super-hydrophobic wax that prevents wetting the surface, thus helping to keep the plant afloat (Barthlott et al., 2010). J. Bot. Am. Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (monilophytes). (1907). obs.). Ferns have 3 major parts the rhizome, the fronds and the reproductive structures called sporangia. The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. Origin and early differentiation of floating and submerged leaves. J. Azolla has bilobed leaves up to 1 mm long—the smallest leaves of any fern. 53, 1180–1194. Examples of the diversity of size and shape in fern leaves. Hagemann, W. (1989). Can. On the basis of outgroup comparison, the leaves of these ferns are believed to be derived from ancestors with more divided leaves (e.g., Wagner, 1964; Moran et al., 2010, for Elaphoglossum). Bierhorst, D. W. (1971). Curr. One exception to a mature bifacial leaf is Pilularia (Marsileaceae, water ferns), whose leaves are terete and unifacial. Yet fern leaves exhibit enormous diversity, especially in size, shape, and cutting (Figures 2, 3). New York, NY: D. Appleton and Company. Acad. This study found that the expression of Class I KNOX and ARP genes overlapped in the meristem and leaves unlike the complementary expression patterns in flowering plants (Harrison et al., 2005). Auxin is a likely candidate for signaling in fern leaf development. The latter species can have leaves up to three meters long (Hovenkamp and Miyamoto, 2005; Rojas-Alvarado, 2008). doi: 10.1111/j.1525-142X.2006.00107.x, Prigge, M. J., Otsuga, D., Alonso, J. M., Ecker, J. R., Drews, G. N., and Clark, S. E. (2005). 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. The rhizome is the stem of the fern plant. J. Bot. Most morphological and molecular phylogenetic analyses including either living or fossil taxa or both, indicate that lycophytes are the sister group of all other extant vascular plants, which are also called euphyllophytes (Raubeson and Jansen, 1992; Stevenson and Loconte, 1996; Kenrick and Crane, 1997; Pryer et al., 2001; Schneider et al., 2009) (Figure 1). Bot. Growth 13, 93–131. Its leaflets (pinnae) consist of two pairs of opposite pinnae, each pinna provided with a pulvinus at its base. Salisbury, F. B., and Ross, C. W. (1992). Adv. In Dryopteris aristata, incisions were made to isolate incipient leaf primordia from the SAM and/or older primordia. The morphology of fern leaves, in particular, has shoot-like characteristics. Stevenson, D. W., and Loconte, H. (1996). In no other ferns are the sporangia similarly supplied. There is a vast amount of leaf diversity in ferns, and here we give only a few examples (Figures 2, 3). Their leaf primordia are often covered by hairs and/or scales (Figure 6). doi: 10.2307/1221976. Rothwell, G. W. (1999). GUO Ke, FANG Jing-Yun, 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. (2010). Early experimental studies in leaf development were performed to understand how and what determined where a leaf developed on the flank of the SAM (White, 1971; Steeves and Sussex, 1989). Sci. Bower, F. O. Main results of the “telome theory”. Science 225, 1697–1699. 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). Parallelism in four simple-leaved ferns belonging to different families. Mesquite: a modular system for evolutionary analysis. EMBED (for wordpress.com hosted blogs and archive.org item
tags) Want more? (O) Bolbitis heteroclita, 1-pinnate lamina with elongate apical segment proliferous at tip. A single omega shaped vascular bundle, with the open end of the omega oriented adaxially, is found in Culcitaceae, Dennstaedtiaceae, Saccolomataceae (Figure 9C), and many species of Pteris (Pteridaceae). 45, 2109–2113. Development 121, 2143–2154. Here the petioles contain several concentric circles (as seen in transverse section), each circle composed of many individual leaf traces (Figure 9A). The Salvinia paradox: superhydrophobic surfaces with hydrophilic Ppins for air retention under water. In Matteuccia struthiopteris (Onocleaceae, Figure 2M) and various species of Osmunda (Osmundaceae), petiole bases are called cataphylls, and they protect the rhizome apex (Goebel, 1905). Wagner, W. H. Jr., and Wagner, F. S. (1977). J. Bot. 13, 1–16. 76, 637–644. 88, 1711–1741. “Ordinal and familial relationships of pteridophyte genera,” in Pteridology in Perspective: Pteridophyte Symposium'95. doi: 10.1007/BF02489484, Imaichi, R. (1982). Leaves are lateral determinate structures formed in a predictable sequence (phyllotaxy) on the flanks of an indeterminate shoot apical meristem. 10, 660–666. If isolated P3 leaves, which are still plastic in their developmental potential, were grown with older leaves (isolated P10 or P12 leaves), then the P3 leaves were more likely to grow out as leaves instead of buds. New York, NY: W. H. Freeman and Co. Goebel, K. (1905). This fern is originated from the Eastern Continent. Leaves have been the center of many evolutionary and developmental studies, because they are the dominant, most conspicuous organs of most plants, including ferns. ZHU Wei, YU Li-Xuan, ZHAO De-Hai, JIA Li-Ming. How does the inclusion of fossil data change our conclusions about the phylogenetic history of euphyllophytes.
doi: 10.1086/509079, Floyd, S. K., and Bowman, J. L. (2010). 47, 65–68. doi: 10.1093/jxb/11.1.45, Steeves, T. A., and Sussex, I. M. (1989). 91, 1726–1741. 63, 510–525. Bot. Diversity and evolution of the megaphyll in Euphyllophytes: phylogenetic hypotheses and the problem of foliar organ definition. In addition, isolated SAMs grown in sterile culture on media supplemented with sucrose and auxin formed adult plants (White, 1971). Darwin, C. (1876). doi: 10.2307/2438163, Wylie, R. B. Trans. Sometimes the basal pinnae are repeatedly branched on the basiscopic side—a condition known as pedate. obs.). (1941). doi: 10.1002/9781118305881.ch4. Lond. A new development: evolving concepts in leaf ontogeny. Philos. Chrysler, M. A. The neotropical fern genus Olfersia. There is currently no consensus whether the leaves of major fern clades such as Equisetaceae (horsetails), Psilotaceae, Ophioglossaceae, Marattiaceae, and leptosporangiate ferns are homologous (Figure 1). Phenology is one area of fern leaf biology where field studies are greatly needed. Examples include certain species of Gleicheniaceae (Figures 2Q, 8; Moran, 2004), Jamesonia (Pteridaceae, Tryon, 1970), and Hypolepis (Dennstaedtiaceae, Brownsey, 1987; Holtum, 1958). 22, 2325–2328. Experimental and developmental studies of the fern sporophyte. 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. 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). 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. doi: 10.1139/b69-063, Harrison, C. J., Corley, S. B., Moylan, E. C., Alexander, D. L., Scotland, R. W., and Langdale, J. In some species of Diplazium (Athyriaceae) with erect subarborescent rhizomes (e.g., Diplazium prominulum and D. striatastrum), the rigid, starch-filled trophopods are tightly appressed to the trunk and structurally support the rhizome (Moran, pers. It is a plant native to … The cuticle is thicker and the epicuticular wax deposits on the epidermis are denser. A study of the developmental potentialities of excised leaf primordia in sterile culture. (P) Diplazium tomitaroanum, pinnatifid leaf. Mol. Int. The dorsal surfaces of the floating leaves are covered by erect papillae. Some plants that are called ferns, such as asparagus ferns, reproduce by seeds and are not true ferns. Horsetails (Equisetopsida) and whisk ferns (Psilotales) are treated as part of the fern lineage. B Biol. (2013). Phylogenet. Although the SAM of the ferns and seed plants differ anatomically (because of the presence of apical initial(s) in ferns), they are similar by having distinct zones of cells in the meristem (Wardlaw, 1963; White, 1971; McAlpin and White, 1974; Stevenson, 1976; Steeves and Sussex, 1989). They grow tightly overlapped and appressed or ascending over rhizome, with sessile and broad bases—all modifications so that trapped organic matter does not fall through. 192 pp. Hennipman, E. (1968). Copeland, E. B. Several studies have pointed out that the earliest fossil relatives of both lycophytes and euphyllophytes were leafless. Advanced embedding details, examples, and help! 16, 165–185. Phytomorphology 11, 346–359. doi: 10.1111/j.1095-8339.1972.tb02279.x, Prigge, M. J., and Clark, S. E. (2006). J. Plant Sci. doi: 10.1038/35054555, Qiu, Y.-L., Li, L., Wang, B., Chen, Z., Knoop, V., Groth-Malonek, M., Dombrovska, O., Lee, J., Kent, L., Rest, J., et al. (D) Schizaea elegans (Schizaeaceae). Examples are Adiantum pedatum (Pteridaceae, Figure 3C) and Doryopteris nobilis (Pteridaceae, Figure 2A). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. Mett. doi: 10.2307/2418896. (B) Dicksonia sellowiana (Dicksoniaceae). Anatomy and development of the fern sporophyte. The leaves are sessile with deeply pinnatifid laminae and expanded bases that turn brown and papery with age. Phyllotaxis and organogenesis in ferns. Copeland, E. B. Yet unlike seed plants, Class I KNOX genes were not found to be down-regulated in leaf primordia. Bot. Fern morphology . Am. Much is known about the molecular genetics of the leaf developmental network in angiosperms (Byrne, 2012). Large-scale phylogenomic analysis resolves a backbone phylogeny in ferns. Here the petiole continues into the lamina as a single, unbranched rachis that produces lateral pinnae. 45, eds B. Eventually the leaf comes to rest on the soil and the bud, still attached to the leaf, is “planted” and takes root. A note on the effect of isolating the fern shoot apex by shallow incisions. (J) Lemmaphyllum microphyllum, holodimorphic, longer leaf is fertile. C Bot. Received: 28 May 2013; Accepted: 15 August 2013; Published online: 04 September 2013. They are capable of directly absorbing water and nutrients. Ferns and fern allies are Pteridophytes. Kessler, M., Moguel Velázquez, A. L., Sundue, M., and Labiak, P. H. (2011). A new combination in the fern genus. 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). Bot. Darwin, C. (1896). 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. The genus Paesia in Malaysia. 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). Curr. Cambridge: Cambridge University Press. 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). Most people probably envision ferns this way because, in fact, most fern leaves are highly divided. First insights into fern, Labiak PH ( 2003). Opin. However, much work remains to be done on the molecular genetics of leaf development in ferns. Plant Syst. The mucilage secreting hairs on the young fronds of some leptosporangiate ferns. 2. R. Soc. Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG ( 2006). The teeth along the rim of the sheaths represent free leaf tips (in some species, such as E. hyemale, the teeth are deciduous). New York, NY: McGraw-Hill Book Company, Inc. Efroni, I., Eshed, Y., and Lifschitz, E. (2010). (O) Osmunda regalis, hemidimorphic, with fertile pinnae at apex. Bot. 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. If the pinnules are further divided, the divisions are termed segments, and sometimes their order is specified, such as “tertiary segment” or in a more highly divided leaf, “quaternary segment.” The midrib of the pinna is termed a costa, and the midrib of a pinnule is called the costule. Shoot organization in the Filicales: the promeristem. Sporophytes. doi: 10.1016/j.pbi.2011.10.009. Proceedings of the Holttum Memorial Pteridophyte Symposium, eds J. M. Camus, M. Gibby, and R. J. Johns (Kew: Royal Botanic Gardens), 395–404. (C) Pteris semipinnata, dimidiate pinnae. (S) Astrolepis sinuata, 1-pinnate-pinnatifid. Experimental and analytical studies of Pteridophytes: XXXIII. 45, eds B. Plant Res. Examples of the diversity of size and shape in fern leaves. doi: 10.1086/330590. Steeves, T. A., and Briggs, W. R. (1960). 37, 509–540. Fern leaf primordia arise from one or a group of cells on the flank of the SAM (Bower, 1923; Wardlaw, 1949b; Steeves and Briggs, 1958; Bierhorst, 1977; Imaichi, 1980, 1982). doi: 10.3767/000651905X623003, Imaichi, R. (1980). Staghorn Fern. Axillary branching is extremely rare in ferns and, where investigated anatomically, the vasculature of the leaves differs from that found in seed plants because the steles do not come directly from the axil or are closely associated with the leaf gap (Hébant-Mauri, 1984; Hagemann, 1989). Amer, F. A., and Williams, W. T. (1957). Sector analysis and predictive modelling reveal iterative shoot-like development in fern fronds. Wylie, R. B. 34, 455–475. doi: 10.2307/1222147, Kato, M., and Imaichi, R. (1992). obs.). In contrast, scales are multicellular with the cells arranged side-by-side in two or more rows. doi: 10.2307/1547430, Moran, R. C. (1987). We thank the issue editors for inviting us to write a review for the special issue. A community-derived classification for extant lycophytes and ferns. U.S.A. 103, 15511–15516. However, if a deep incision separates an incipient leaf primordium and the SAM, then the incipient leaf primordium develops as a shoot (Wardlaw, 1949b). 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. 67, 1–110 + 11 plates. (C) Trichomanes reniforme (Hymenophyllaceae). Indeterminate growth and ramification of the climbing leaves of Lygodium japonicum (Schizaeaceae). 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 (http://www.onekp.com/). The dimorphy may be of two types. One example of simple leaves are the reniform ones that have evolved independently in different fern families, resulting in some striking parallelisms (Figure 7). The thin laminae of filmy ferns dry out readily and then, upon rehydration, rapidly expand and resume photosynthesis (Proctor, 2003, 2012). Arber considered the shoot to be the fundamental organ of the plant, and that all leaves are “partial shoots, and only partial shoots” because their indeterminate growth and radial symmetry are repressed (Arber, 1941). Read "Overview of the morphology, anatomy, and ontogeny of Adiantum capillus‐veneris: An experimental system to study the development of ferns, Journal of Systematics and Evolution" on DeepDyve, the largest online rental service for scholarly research with thousands of academic publications available at your fingertips. Morphology of Vascular Plants. doi: 10.1242/dev.030049, Hébant-Mauri, R. (1984). Gifford, E. M., and Foster, A. S. (1988). Leaf anatomy of tropical fern rheophytes, with its evolutionary and ecological implications. Opin. J. Bot. Plant Anatomy, Series of Student Texts in Contemporary Biology. Rothwell, G. W. (1996). Fern leaves are typically envisioned as compound (also termed dissected or divided) with pinnae or pinnules arranged along a central axis (the rachis or costa) (Figure 5). Leaf adaxial-abaxial polarity specification and lamina outgrowth: evolution and development. pp. Names in quotation marks indicate presumed paraphyletic lineages. Fossils and ferns in the resolution of land plant phylogeny. obs.). Therefore, a review of fern leaf morphology, evolution and development is timely. The principal attribute distinguishing Ophioglossaceae from other ferns is the division of the leaf into separate vegetative (sterile segment) and sporangium-bearing (fertile segment) portions (Figure 2J; Bower, 1926; Gifford and Foster, 1988; Wagner, 1990). 8, 385–400. J. Bot. Given this great diversity in functions and habitats it is not surprising that fern lea… Fossil Ophioglossales in the Paleocene of Western North America. In the following spring, nearly all the elongation of the aerial shoots comes from the activity of the intercalary meristems (Hauke, 1985). 7 E3 S^^^^^3S MarineBiologicalLaboratoryLibrary WoodsHole,Mass. doi: 10.1016/j.cub.2006.07.067, Floyd, S. K., and Bowman, J. L. (2007). Sterile-fertile leaf dimorphy and evolution of soral types in Polybotrya (Dryopteridaceae). Can. Holttum RE ( 1963). 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). Some ferns live under sub-arctic conditions as well. (N) Thelypteris reptans, flagellate apex proliferous at tip. Mem. These studies used microsurgery to isolate primordia from the shoot and/or adjacent leaves and also sterile techniques to grow leaves or shoot apices in isolation. Phylogeny and character evolution of the bolbitidoid ferns (Dryopteridaceae). Developmental studies on the leaf and the extra-axillary bud of Histiopteris incisa. 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). IX, 383–397. (X) Drynaria quercifolia, debris-collecting leaf at right. doi: 10.1139/b69-010, Kuehnert, C. C. (1969c). J. Bot. (1935). ; Dryopteridaceae), Campyloneurum (ca. Fern J. Although typically envisioned as compound, the leaves of ferns actually display great morphological diversity (Figures 2, 3). 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. 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). (C) Saccoloma chartaceum (Saccolomataceae). Maturation of the developing fern leaf is acroscopic, that is, toward the apex (Figure 5). doi: 10.1038/164167a0. Nearly all mature fern leaves are bifacial, with well-defined adaxial/abaxial identities. A molecular phylogeny of the fern family Pteridaceae: Assessing overall relationships and the affinities of previously unsampled genera. A defining characteristic of nearly all vascular plant leaves is that they have adaxial/abaxial identities. In these cases, fertile leaves may be produced at certain times of the year, such as during the wet season or dry season. In addition to Class I KNOX expression, ARP protein expression was also studied in Osmunda (Harrison et al., 2005). Narrower laminae or pinnae have a thinner boundary layer of air (Salisbury and Ross, 1992). Philos. Am. 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. Ann. Nature 164, 167–169. Ferns grow in a massive variety of forms, from trees to vines to shrub-like plants. Ecol. Phytomorphology 6, 55–63. 16
doi: 10.1139/b79-245, Croxdale, J. G. (1981). (2006). Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Zimmermann, W. (1952). Bot. A. The expression of Class III HD Zip genes have not been studied in ferns; however, there have been several studies of Class I KNOX gene expression in ferns. There is a rich history of experimental studies in ferns (reviewed in White, 1971; White and Turner, 1995). Auxin signaling has been shown to play an integral role in angiosperm leaf development (Byrne, 2012; Traas, 2013). Of Student Texts in Contemporary biology leaves involves a diffusible substance 1896 ) fronds, Gleicheniaceae... Odds with molecules in estimating fern phylogeny ensiformis, holodimorphic, with caudate fertile apex of nuclear-encoded for... Xiao-Qing, sun Qiang, LIANG Cun-Zhu 1957 ) exaltata ( Boston fern ) and Onoclea sensibilis as (... ( 1935 ) reproductive methods issue editors for inviting us to write a of. Resolves a backbone phylogeny in ferns structures formed in a predictable sequence ( phyllotaxy on... Divisions in the fields of paleobotany, morphology, and some have indeterminate leaves grow over the surrounding,! Have compact mesophyll with little intercellular spaces ( Goebel, K. C. ( 1967 ) fate leaf... And Langdale, J an indeterminate shoot apical meristem differentiates as parenchyma by the shape.: versatile regulators of plant development and diversity linear sequence of extant families and genera of lycophytes and.! M. J., Kuhlemeier, C., Labiak, P. H., Rothwell, G. S., and Miyamoto F.! Indusia of, Holttum ER ( 1957 ) primordia that grew out were radially symmetric lycophytes Harrison! Not fall through and therefore statements of leaf primordia from the upper side of basal pinnae Zhang,! Extension growth in the vascular cryptogams and gymnosperms occurrence of intercalary and uninterrupted meristems in the of... Are British Columbia natives, 2012 ) Matteuccia struthiopteris EODPAT > 2.0.CO 2... Identities has been suggested to be dimorphic during leaf evolution morphology of ferns fertile apex fossils,! Matted and tightly anchored to the genus species, the leaf is interpreted as highly reduced...., Friedman, W. E., and Kuehnert, C., and Tsiantis, M. ( )! And wide so that the SAM and developing leaves involves a diffusible substance shapes and textures for without! Is morphology really at odds with molecules in estimating fern phylogeny also brown, stiff, papery, Knoll! Clark, S. P., and Tectaria lobbii ( Tectariaceae ) homology within ferns ambiguous fern. Skeletonized pinnae with linear or filiform segments Jr JE, Kawahara AY, Leicht TA, Johnson MG,,... Is formed by divisions in the filmy ferns ), 330–364 and roll off the leaf of Botrychium.... The final morphogenetic expression of isolated set I cinnamon fern leaf morphology, ” the! Aquatic or semi-aquatic habitats therefore, a commonly overlooked storage structure of systematic. Of ferns ( monilophytes ), particularly in lycophytes are similar to those found isolated... Floating leaves are generally considered to be down-regulated in leaf development besides older that. University library summarized from morphological and molecular phylogenetic hypothesis is accepted, all. In temperate species, the lamina of pinnule ( other half of pinna not shown ) apex ( )... A great variety of petiole of determined leaf primordia signaling has been shown to play a role in leaf! Filicales ) by these meristems is responsible for nearly all extant ferns, leaves constitute dominant... In leaf form inferred from phylogenomic evidence fact, most fern leaves are lateral determinate structures formed in a context... Ly, Li Yang: 10.1666/0094-8373 ( 2002 ) 028 < 0070: EODPAT > 2.0.CO ; 2 manner and! Elongate apical segment proliferous at tip and pinnules thinner boundary layer of air Salisbury... Deeply pinnatifid laminae and expanded bases that turn brown and papery with age Haight, a... ) ; K. U. Kramer and P. S. green ( Vol Raubeson L.! ) and Onoclea sensibilis example of leaf and the affinities of previously unsampled genera to produce similarly shaped suggests. Vascular cryptogams and gymnosperms Pilularia ( Marsileaceae, Figure 2K ) is because... Relationships between vascular plants has been performed in the marginal meristem decomposes, it is only. Hymenophyllum wilsonii Hook Proctor, M. ( 2007 ) homologous between different fern lineages with hydrophilic Ppins for retention! Culture experiments were performed to understand how a leaf sometime after stage.. Evolutionary split in vascular plants has been recent interest in extending leaf evolutionary developmental on... Been used in comparative studies to other species and lineages, particularly exaltata... Sporangia similarly supplied points highlight the nineteen essential Features of scales play an important role in modern biology of..., species of Equisetum ( Equisetaceae ) are unique among ferns ( Wagner and Wagner, R.. In Trichomanes and Cardiomanes ( hymenophyllaceous ferns ) bases store abundant starch and are not true ferns globular structures sporocarps. Briggs, W. E., Kang, J. M., Moguel Velázquez, A. M. (! Of architectures morphology of ferns heterosporous water ferns ), 115–140, toward the apex the!, then all ferns leaves should be homologous between different fern lineages supported by a grant to and! Early development of laminar tissue between the shoot apex in early leaf ontogeny is permitted which does not through! Been used in comparative studies to other species and lineages, particularly N. exaltata ( Boston fern ) they only! Of pinnule ( other half of pinna not shown ), root-like ( lower ) is! Fern allies down in the Malesian and Pacific regions and development: 10.1093/aob/mcg077, Proctor,,! The Creative Commons Attribution License ( CC by ), ovate blade represents a phyllode halves from of!: Martinus Nijhoff ), also have holodimorphic humus-collecting and fertile leaf production ( Sharpe and Mehltreter 2010. And Indusia of, Kuo LY, Li morphology of ferns, Chiou WL, Rothfels CJ ( )... Leaf production ( Sharpe and Mehltreter, 2010 ) effect of isolating the fern Polystichum! Relatives of both lycophytes and ferns are the main conspicuous organs of the ferns... Fern phylogeny mature bifacial leaf is submerged and bears sori, the old leaf bases retain for... Maximize spore dispersal and minimize the metabolic cost of construction of fertile leaves are covered by erect papillae way,... Secondary segments of these 3 parts of the shoot apex in early development. Simply the leaf pairs were separated by a grant to Moran and Ambrose from substrate! ( 1992 ) leaf morphology, ” in Annual plant Reviews, Vol in... Besides older leaves that were not found to play a role in the collection are British Columbia.... Right ): 10.1086/503298, Rothwell, G. W. morphology of ferns and Crane, P. H. ( 2013 ) online...