doi: 10.1242/dev.030049, Hébant-Mauri, R. (1984). (1926). Finally, the frequency of occurrence of stomata per unit area of leaf is higher in rheophytes than in related upland species (Kato and Imaichi, 1992). doi: 10.5962/bhl.title.26129. Fern leaves generally exhibit finite (determinate) growth but with longer meristematic activity of the apical portion and maturation toward the apex (acroscopic growth). Steeves, T. A., and Briggs, W. R. (1960). 192 pp. J. In addition, studies of leaf development genes in Psilotum (Psilotaceae), Ophioglossum (Ophioglossaceae), and Equisetum (Equisetaceae) could provide crucial molecular data to better interpret the leaf morphology of these distinctive genera. doi: 10.2307/2438163, Wylie, R. B. 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. Philos. 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). doi: 10.1086/330590. A KNOX family TALE. In other ferns, the leaf bases store abundant starch and are termed trophopods (Wagner and Johnson, 1983). Ferns and fern relatives are known as Pteridophytes. Byrne, M. (2012). Experimental and analytical studies of Pteridophytes: XXXIII. Experimental studies of fern leaves have investigated the growth of heteroblastic leaf sequences, cataphylls, and simple and compound leaves (Wardlaw, 1963; White, 1971). 200, 54–174. Some are twining and vinelike; others float on the surface of ponds. The first kind of leaf, of which there are two in the false whorl, is floating, green, entire, and conduplicate in bud (i.e., not circinate). Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. (1952). doi: 10.1007/BF00708854. Imaichi, R. (2008). This condition is shared with the lycophytes (Moran, pers. Can. (1907). A., Hasebe, M., and Juarez, C. M. (2005). Cambridge: Cambridge University Press. The spores are bilateral with a smooth exine and the perine smooth in Cyrtomium and granulose to … Unlike seed plants, ferns and their allies reproduce by spores. The thin laminae of filmy ferns dry out readily and then, upon rehydration, rapidly expand and resume photosynthesis (Proctor, 2003, 2012). Plant Cell Physiol. Narrower laminae or pinnae have a thinner boundary layer of air (Salisbury and Ross, 1992). Club 21, 86–95. (1936). J. However, Wardlaw (1949b,d) concluded that initially leaf primordia and shoot buds of Dryopteris aristata are histologically indistinguishable. 91, 717–727. 中国产瓦韦属植物的分类学研究. Morphology and evolution of epiphytic Davalliaceae scales. Rhythmic leaf extension growth in the Gleicheniaceae (from Moran, 2004). Ferns are flowerless green plants . Flora of China,Vol. Gazette 52, 151–153. Lond. The leaves are highly reduced and connate laterally to form a sheath around the base of each segment of the aerial stem. 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. That half of a pinna or pinnule that occurs on the side toward the distal apex of the axis that bears it is called the acroscopic side, and accordingly, the side that occurs toward the base is the basiscopic (for more details of fern leaf terminology, see Tryon, 1960). Int. Alphen aan den Rijn: Sijthoff and Noordhoff. Furthermore, ferns grow in many habitats—from mangroves at sea level to alpine vegetation above tree line, temperate forests to arctic tundras, and desserts to wetlands. The 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. 61, 562–579. This fern is originated from the Eastern Continent. The teeth along the rim of the sheaths represent free leaf tips (in some species, such as E. hyemale, the teeth are deciduous). R. Soc. Ferns General Characteristics. Scrambling ferns often have leaf apices exhibiting intermittent growth (Figure 8). A monograph of the fern genus Platycerium (Polypodiaceae). Towards resolving the complete fern tree of life. The condition is also present anatomically two ways: first, with the mesophyll differentiated adaxially as palisade tissue and abaxially as spongy tissue, and second, by greater elongation of cells on the abaxial side vs. adaxial. Plant Sci. Molecular genetic studies in angiosperm leaf development are beginning to provide clues into the nature of the signaling that occurs between the shoot apex and leaf primordia (Byrne, 2012). Simple leaves in ferns are found in Elaphoglossum (ca. The SAM in ferns typically has 1 or 2 large apical cells surrounded by small cytoplasmically dense cells that divide frequently (Bower, 1884; Wardlaw, 1963; Bierhorst, 1971; White, 1971; McAlpin and White, 1974; Stevenson, 1976; White and Turner, 1995). In Equisetum, the relationship of the vegetative and fertile portions has been debated: the sporangiophore (the fertile portion) has been interpreted as being a novel organ “(organ sui generis)” or homologous to the leaves (Goebel, 1905; Bower, 1935; Zimmermann, 1952; Page, 1972). Oxford: Clarendon Press. In these experiments the leaf pairs were separated by either an impermeable barrier or a permeable barrier. Morphogenesis of simple and compound leaves: a critical review. Most ferns are leptosporangiate ferns… Furthermore, species of Ophioglossaceae usually produce only one leaf at a time. Salvinia leaves. ; Polypodiaceae), and Vittaria (6 spp. (B) Deparia acrostichoides, lamina 1-pinnate-pinnatifid. Class I KNOX and Class III HD Zips genes have been well-studied across the flowering plants and to a lesser extent in gymnosperms (reviewed in Floyd and Bowman, 2007; Efroni et al., 2010; Floyd and Bowman, 2010; Hay and Tsiantis, 2010; Byrne, 2012; Townsley and Sinha, 2012; Yamaguchi et al., 2012). In the experiments with Osmunda cinnamomea, most of the excised P3 leaves develop as buds; however, some still develop as leaves. Can. 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). Developmental potentialities of leaf primordia of Osmunda cinnamomea. Kaplan, D. R., and Groff, P. A. (U) Hemionitis ariifolia, hastate at left, deltate at right (from same plant). It is a plant native to … The twining results from the widely circumnutating leaf being interrupted by contact with a support, and, when the rope hits a pole, the rope continues its motion around the pole, wrapping upward and thus climbing the pole (Darwin, 1876). Morphology and Anatomy. Yet unlike seed plants, Class I KNOX genes were not found to be down-regulated in leaf primordia. Bot. Ferns have 3 major parts the rhizome, the fronds and the reproductive structures called sporangia. Can. Hagemann, W. (1989). Auxin has been shown to effect the leaf complexity in the fern Marsilea as well as crozier development (Allsopp, 1952; Steeves and Briggs, 1960). Experimental and analytical studies of pteridophytes. R. Soc. J. Bot. 11, 403–418. Ferns are seedless vascular plants of humid tropics and temperate areas. (A) Eupodium laeve (Marattiaceae). One process involved the elongation of some of the branches more than others, producing a main central branch with subordinate lateral ones (overtopping). 47, 65–68. (C) Trichomanes reniforme (Hymenophyllaceae). The dorsal surfaces of the floating leaves are covered by erect papillae. Sec. (D) Acrostichum danaeifolium (Pteridaceae). doi: 10.1666/0094-8373(2002)028<0070:EODPAT>2.0.CO;2. Tweede Reeks 80, 1–126. 70, 682–690. Mem. The rhizome scales of, Kuo LY, Li FW, Chiou WL, Wang CN ( 2011). The character differences maximize spore dispersal and minimize the metabolic cost of construction of fertile leaves (Moran, 1987). In all vascular plants, leaves arise from the SAM. Most people probably envision ferns this way because, in fact, most fern leaves are highly divided. Generally such leaves are either pendulous or scrambling over the surrounding vegetation. (Y) Aglaomorpha meyeniana, hemidimorphic, with narrow distal pinnae fertile and base expanded for collecting fallen organic debris. Rothwell, G. W. (1996). Waites, R., and Hudson, A. J. Bot. 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). J. Bot. Mol. “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. Trends Plant Sci. 56, 1982–1991. Wardlaw, C. W. (1949b). Blumea 16, 97–103. Mater. 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. Daigobo, S. (1972). Experiments on phyllotaxis. Fern leaves and megaphylls of other groups are defined by a combination of characters that are a result of specific developmental processes. Philos. 北京: 科学出版社. 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. (X) Drynaria quercifolia, debris-collecting leaf at right. Molecular phylogenetic hypotheses of extant taxa consider ferns as a monophyletic group that includes the Equisetaceae, Psilotaceae, and Ophioglossaceae (Figure 1; Pryer et al., 2001; Qiu et al., 2006; Grewe et al., 2013). 93, 25–38. Usually growing underground or right along the surface, these horizontal stems hold the vascular system that transports water and nutrients. Part 1, 2nd Edn. ; Polypodiaceae), Microgramma (35 spp. 169-256. doi: 10.1098/rstb.1935.0007. doi: 10.1086/653130. of fern morphology is the presence of epidermal ap-pendages. Univ. Plant Physiology. 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. Sci. doi: 10.2307/1221976. Holttum RE ( 1949). KNOX homeobox genes potentially have similar function in both diploid unicellular and multicellular meristems, but not in haploid meristems. doi: 10.1111/j.1095-8339.1991.tb00220.x, Doyle, J. They are usually easy to recognize by the featherlike shape of their leaves, which are called fronds. Die Phylogenie der Pflanzen; ein Uberblick über Tatsachen und Probleme. Contrib. The scientific name of ostrich fern is Matteuccia struthiopteris. Distinct developmental mechanisms reflect the independent origins of leaves in vascular plants. In many species, a single frond is often divided multiple time along the length of the frond. Class III homeodomain-leucine zipper gene family members have overlapping, antagonistic, and distinct roles in Arabidopsis development. J. Microsurgery experiments have provided further insight into the relationship between the shoot apical meristem and leaf development. The transition from vegetative to reproductive growth of shoot apices of holoheterophyadic species of Equisetum: phenology, morphology, and anatomy. White, R. (1971). Morphology and systematics of the Cycadales. 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. obs.). 76, 161–178. 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). Straits Settlements 30, 251–267. Abstract. These results might suggest either the independent origin of megaphylls in ferns and seed plants, or may simply reflect the prolonged indeterminacy of fern leaves. Figure 9. In: Ambrose BA, Purugganan M, eds. J. Bot. This is a very distinct looking fern, and while it can be grown for some time in a pot … Am. Bot. Being the main conspicuous organ of nearly all vascular plants and often easy to recognize as such, it seems surprising that leaves have had multiple origins. II. In these cases, fertile leaves may be produced at certain times of the year, such as during the wet season or dry season. Light and desiccation responses of some Hymenophyllaceae (filmy ferns) from Trinidad, Venezuela and New Zealand: poikilohydry in a light-limited but low evaporation ecological niche. 15, 24–30. Lond. ZHU Wei, YU Li-Xuan, ZHAO De-Hai, JIA Li-Ming. Zhang GM, Liao WB, Ding MY, Lin YX, Wu ZH, Zhang XC, Dong SY, Prado J, Gilbert MG, Yatskievych G, Ranker TA, Hoope EA, Alverson ER, Metzgar JS, Funston AM, Masuyama S, Kato M ( 2013). Experiments on the cause of dorsiventrality in leaves. In general, leaves are the main conspicuous organs of vascular plants and often easy to recognize as such. The morphology of fern leaves, in particular, has shoot-like characteristics. Brittonia 63, 233–244. A glossary of some terms relating to the fern leaf. Opin. These experiments indicate that a Dryopteris aristata leaf is determined as a leaf sometime after stage P1. Neerl. Phylogenet. Examples are Adiantum pedatum (Pteridaceae, Figure 3C) and Doryopteris nobilis (Pteridaceae, Figure 2A). Rothwell, G. W. (1999). Phytomorphology 8, 60–72. J. Hairs are uniseriate and either one-celled or multicelled. Iowa Acad. Rev. Their leaf primordia are often covered by hairs and/or scales (Figure 6). 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With Osmunda cinnamomea, most of the leaf is represented by either impermeable... Experimental induction of buds from leaf primordia on undetermined leaf primordia in sterile culture on media supplemented sucrose!
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