Ceratopteris

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Ceratopteris
Ceratopteris cornuta in aquarium alone.jpg
Ceratopteris cornuta in aquarium
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Family: Pteridaceae
Subfamily: Parkerioideae
Genus: Ceratopteris
Brongn.
Type species
Ceratopteris thalictroides
Species

See text

Synonyms
  • CryptogenisRichard 1823 ex Brongniart 1823
  • EllebocarpusKaulfuss 1824
  • FurcariaDesvaux 1827
  • ParkeriaHooker 1825
  • TeleozomaBrown 1823

Ceratopteris is the only genus among homosporous ferns that is exclusively aquatic. It is pan-tropical and classified in the Parkerioideae subfamily of the family Pteridaceae. [1]

Contents

Description

Ceratopteris cornuta in detail Ceratopteris cornuta in detail (emerse).jpg
Ceratopteris cornuta in detail

Erect aquatic or subaquatic ferns of moderate size. Rhizome short, fleshy, horizontal and ascending to erect, loosely rooted in the mud or +/- floating, radial, dictyostelic with numerous meristeles and medullary strands, young parts bearing thin, ovate, +/- cordate, clathrate scales. Fronds stipitate, the stipes fleshy, with numerous longitudinal air canals, abaxially rounded and ribbed, adaxially flattened, vascular bundles in a peripheral ring, one with each rib and several to the adaxial side, and several smaller medullary strands; lamina dimorphic, sterile fronds +/- spreading, 2–3-pinnatifid with broad membranous lobes, venation reticulate without included free veinlets, often with proliferous buds in the axils; fertile fronds erect, longer, narrower and more divided than the sterile, the lobes strongly recurved to completely cover the adiaxial surface, venation longitudinal, branching at the bases of the lobes. Sporangia solititary, scattered along the veins, exindusiate but protected by the continuous reflexed margin of the lamina, large, short-stalked, annulus broad, irregular, of 30–70 thickened cells, or lacking, containing 16 to 32 spores. Spores large, trilete, ribbed with irregular long meshes. [2]

Taxonomy

Ceratopteris was long placed in the monogeneric family Parkeriaceae, thought to be unique because of its aquatic adaptations. However, recent genetic analysis has shown it to be clearly allied with Acrostichum in the subfamily Parkerioideae, within the family Pteridaceae.

At one time, some authorities recognized only one species; now some authorities recognize only four species. However, recent work by Masuyama and Watano has suggested that C. thalictroides actually consists of four cryptic species (thalictroides, froesii, gaudichaudii, oblongibloba). [3]

Phylogeny

Nitta et al. 2022 [4] and Fern Tree of life [5]
Ceratopteris

C. richardii Brongn. 1823 ("C-fern"; Triangle waterfern)

C. cornuta (P. Beauv.) Le Prieur 1830

C. chingii Yang & Yu 2022

C. chunii Yang 2022

C. shingii Yan & Zhang 2020

C. pteridoides (Hook.) Hieron. 1905 (Floating antlerfern)

C. thalictroides (L.) Brongn. 1821 (Water Sprite, Oriental water fern)

Other species:

Distribution and habitat

This widespread genus of four to six species inhabits the humid tropics.

Ceratopteris is also important in the study of pteridophytes and is a commonly used model organism for use in genomic studies, due to the ease and rapidity with which it can be grown in laboratories, as well as having well-characterized phenotypes. Patented strains of this plant have been developed.

Outcrossing and self-fertilization

A gametophyte is one of the two alternating multicellular phases of the fern life cycle. The gametophyte is haploid. The other stage is the diploid sporophyte. A gametophyte is a multicellular organism that develops from a haploid spore (that has one set of chromosomes). Ceratopteris thalictroides has two types of gametophytes with different sexual expression: these types are hermaphrodite and male. [6] Hermaphroditic gametophytes have one or several archegonia and a few antheridia. Archegonia are multicellular structures of the gametophyte that produce and contain the ovum or female gamete. The corresponding male organ is the antheridium, a haploid organ producing and containing sperm. The second of the two types of gametophyte are male that produce only antheridia.

When Ceratopteris richardii ferns are grown alone they develop into hermaphrodites (with mainly female archegonia). However, plants that grow near established genetically identical individuals develop into males. The hermaphrodite secretes a pheromone (antheridiogen) that tells others nearby to be male. [7] Thus when plants are near each other outcrossing is promoted. Self-fertilization predominates when there is no nearby partner.

Toxicity

The plants may have carcinogenic properties. [8]

Uses

Ceratopteris is a fairly popular aquarium plant, often sold under the name "water sprite." It may be grown as an emersed but natant (floating) plant, or as an immersed plant rooted in the substrate. Under the right conditions the plants will grow fully emerse erect leaves. Under a bright light the plants can grow fast, and be used to help cycle an aquarium. [9]

This plant is often used as a vegetable, particularly in Asia. [10]

Related Research Articles

<span class="mw-page-title-main">Gametophyte</span> Haploid stage in the life cycle of plants and algae

A gametophyte is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.

<span class="mw-page-title-main">Spore</span> Unit of reproduction adapted for dispersal and survival in unfavorable conditions

In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, fungi and protozoa.

<span class="mw-page-title-main">Alternation of generations</span> Reproductive cycle of plants and algae

Alternation of generations is the predominant type of life cycle in plants and algae. In plants both phases are multicellular: the haploid sexual phase – the gametophyte – alternates with a diploid asexual phase – the sporophyte.

<span class="mw-page-title-main">Bryophyte</span> Terrestrial plants that lack vascular tissue

Bryophytes are a group of land plants, sometimes treated as a taxonomic division, that contains three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. In the strict sense, Bryophyta consists of the mosses only. Bryophytes are characteristically limited in size and prefer moist habitats although they can survive in drier environments. The bryophytes consist of about 20,000 plant species. Bryophytes produce enclosed reproductive structures, but they do not produce flowers or seeds. They reproduce sexually by spores and asexually by fragmentation or the production of gemmae. Though bryophytes were considered a paraphyletic group in recent years, almost all of the most recent phylogenetic evidence supports the monophyly of this group, as originally classified by Wilhelm Schimper in 1879. The term bryophyte comes from Ancient Greek βρύον (brúon) 'tree moss, liverwort', and φυτόν (phutón) 'plant'.

<span class="mw-page-title-main">Sporophyte</span> Diploid multicellular stage in the life cycle of a plant or alga

A sporophyte is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.

<i>Gymnocarpium dryopteris</i> Species of fern

Gymnocarpium dryopteris, the western oakfern, common oak fern, oak fern, or northern oak fern, is a deciduous fern of the family Cystopteridaceae. It is widespread across much of North America and Eurasia. It has been found in Canada, the United States, Greenland, China, Japan, Korea, Russia, and most of Europe. It is a seedless, vascular plant that reproduces via spores and have a life cycle with alternating, free-living sporophyte and gametophyte phases.

<span class="mw-page-title-main">Pteridaceae</span> Family of ferns

Pteridaceae is a family of ferns in the order Polypodiales, including some 1150 known species in ca 45 genera, divided over five subfamilies. The family includes four groups of genera that are sometimes recognized as separate families: the adiantoid, cheilanthoid, pteridoid, and hemionitidoid ferns. Relationships among these groups remain unclear, and although some recent genetic analyses of the Pteridales suggest that neither the family Pteridaceae nor the major groups within it are all monophyletic, as yet these analyses are insufficiently comprehensive and robust to provide good support for a revision of the order at the family level.

Monoicy is a sexual system in haploid plants where both sperm and eggs are produced on the same gametophyte, in contrast with dioicy, where each gametophyte produces only sperm or eggs but never both. Both monoicous and dioicous gametophytes produce gametes in gametangia by mitosis rather than meiosis, so that sperm and eggs are genetically identical with their parent gametophyte.

<span class="mw-page-title-main">Prothallus</span> Gametophyte stage in the fern life cycle

A prothallus, or prothallium, is usually the gametophyte stage in the life of a fern or other pteridophyte. Occasionally the term is also used to describe the young gametophyte of a liverwort or peat moss as well. In lichens it refers to the region of the thallus that is free of algae.

Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from either parent. Asexual reproduction produces new individuals without the fusion of gametes, resulting in clonal plants that are genetically identical to the parent plant and each other, unless mutations occur.

<i>Ceratopteris thalictroides</i> Species of aquatic plant

Ceratopteris thalictroides is a fern species belonging to the genus Ceratopteris, one of only two genera of the subfamily Parkerioideae of the family Pteridaceae.

<span class="mw-page-title-main">Environmental sex determination</span> Method of sex-determination

Environmental sex determination is the establishment of sex by a non-genetic cue, such as nutrient availability, experienced within a discrete period after fertilization. Environmental factors which often influence sex determination during development or sexual maturation include light intensity and photoperiod, temperature, nutrient availability, and pheromones emitted by surrounding plants or animals. This is in contrast to genotypic sex determination, which establishes sex at fertilization by genetic factors such as sex chromosomes. Under true environmental sex determination, once sex is determined, it is fixed and cannot be switched again. Environmental sex determination is different from some forms of sequential hermaphroditism in which the sex is determined flexibly after fertilization throughout the organism’s life.

<i>Lophosoria quadripinnata</i> Species of fern

Lophosoria quadripinnata(J.F.Gmel.) C.Chr. is a species of fern that, according to DNA molecular analysis, belongs to the family Dicksoniaceae, where it is placed in the genus Lophosoria. It is found in the Americas spanning from Cuba and Mexico to Chile. In Chile it is present in the area between Talca and Aysén including Juan Fernández Islands. In Argentina it grows only in the humid valleys of western Neuquén and Río Negro Province. Diamondleaf fern is a common name. In Spanish it is known as 'ampe' or palmilla, but one has to remember that there are several species of ferns called "palmillas" that have larger or smaller fronds, and which grow in colder climates. It is a medium-sized plant, growing to about 4–5 feet and even though the rhizome does not grow a trunk, it is clearly related to the other tree ferns due to features that were apparently already present in their common ancestor, like 'pneumathodes', and the rhizome which changed from the dorsiventral symmetry typical of the other ferns, to a radial symmetry typical of tree ferns. Their large and multiple pinnate fronds, with the petiole raised adaxially, and the hairs on the rhizome and lower part of the petioles, also resemble those of tree ferns. To identify the species, use the position and characteristics of the spores found on the fertile fronds. The genus already existed in the Cretaceous Period in southern Gondwana according to fossil remains found in Antarctica. The species is well known as an ornamental plant.

<span class="mw-page-title-main">Sexual reproduction</span> Reproduction process that creates a new organism by combining the genetic material of two organisms

Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes combines with another gamete to produce a zygote that develops into an organism composed of cells with two sets of chromosomes (diploid). This is typical in animals, though the number of chromosome sets and how that number changes in sexual reproduction varies, especially among plants, fungi, and other eukaryotes.

<i>Ceratopteris richardii</i> Species of aquatic plant

Ceratopteris richardii is a fern species belonging to the genus Ceratopteris, one of only two genera of the subfamily Parkerioideae of the family Pteridaceae. It is one of several genera of ferns adapted to an aquatic existence. C. richardii was previously regarded as being part of the species Ceratopteris thalictroides.

<i>Acrostichum</i> Genus of ferns

Acrostichum is a fern genus in the Parkerioideae subfamily of the Pteridaceae. It was one of the original pteridophyte genera delineated by Linnaeus. It was originally drawn very broadly, including all ferns that had sori apparently "acrostichoid", or distributed in a uniform mass across the back of the frond, rather than organized in discrete sori. This led Linnaeus to include such species as Asplenium platyneuron in the genus, because the specimen he received had sori so crowded that it appeared acrostichoid.

<i>Hymenophyllum australe</i> Species of fern

Hymenophyllum australe, commonly known as austral filmy fern, is a relatively large rupestral and epiphytic fern, indigenous to eastern Australia and New Zealand. It belongs to the unique Hymenophyllum genus, which are characterised by their thin membranous fronds that are seldom more than one cell thick, with the exception of regions over and around veins. Hymenophyllum australe is distinctive in that the fronds are typically thicker than other Hymenophyllum species, often being up to 2-3 cells thick.

<i>Asterotheca</i> Genus of plants

Asterotheca is a genus of seedless, spore-bearing, vascularized ferns dating from the Carboniferous of the Paleozoic to the Triassic of the Mesozoic.

<i>Polyphlebium venosum</i> Species of fern

Polyphlebium venosum, the veined bristle-fern or bristle filmy fern, is a fern in the family Hymenophyllaceae. It is only found in wet forests, mainly growing as an epiphyte on the shady side of the soft tree fern, Dicksonia antartica. It also grows on logs, trunks of trees and rarely on trunks of Cyathea species or on wet rock-faces. It is found in the wetter parts of Eastern Australia and New Zealand. P. venosum has poor long-distance dispersal compared to other ferns due to its short lived spore. Notable features of Polyphlebium venosum include it being one cell layer thick, 5–15 cm in length, having many branching veins and a trumpet shaped indusium.

Antheridiogens are a class of chemicals secreted by fern gametophytes that have "been shown to influence production of male gametangia and thus mating systems in a large number of terrestrial fern species". Antheridiogens are only observed in homosporous fern species, as all gametophytes are potentially bisexual.

References

  1. PPG I (2016), "A community-derived classification for extant lycophytes and ferns", Journal of Systematics and Evolution, 54 (6): 563–603, doi: 10.1111/jse.12229 , S2CID   39980610
  2. http://www.anbg.gov.au/fern/aquatic/parkeriaceae.html Archived 2007-11-05 at the Wayback Machine Ceratopteris in Australian National Herbarium
  3. Masuyama, Shigeo; Yasuyuki Watano (2010). "Cryptic Species in the Fern Ceratopteris thalictroides (L.) Brongn. (Parkeriaceae). IV. Taxonomic Revision". Acta Phytotax. Geobot. 61 (2): 75–86.
  4. Nitta, Joel H.; Schuettpelz, Eric; Ramírez-Barahona, Santiago; Iwasaki, Wataru; et al. (2022). "An Open and Continuously Updated Fern Tree of Life". Frontiers in Plant Science. 13: 909768. doi: 10.3389/fpls.2022.909768 . PMC   9449725 . PMID   36092417.
  5. "Tree viewer: interactive visualization of FTOL". FTOL v1.5.0 [GenBank release 256]. 2023. Retrieved 17 August 2023.
  6. Chen, X.; Chen, Z.; Huang, W.; Fu, H.; Wang, Q.; Wang, Y.; Cao, J. (2019). "Proteomic analysis of gametophytic sex expression in the fern Ceratopteris thalictroides". PLOS ONE. 14 (8): e0221470. Bibcode:2019PLoSO..1421470C. doi: 10.1371/journal.pone.0221470 . PMC   6699692 . PMID   31425560.
  7. Banks, J. A. (1994). "Sex-determining genes in the homosporous fern Ceratopteris". Development. 120 (7): 1949–1958. doi:10.1242/dev.120.7.1949. PMID   7925000.
  8. The Complete Guide to Edible Wild Plants. United States Department of the Army. New York: Skyhorse Publishing. 2009. p. 109. ISBN   978-1-60239-692-0. OCLC   277203364.{{cite book}}: CS1 maint: others (link)
  9. "How to Plant & Grow Water Sprite (Ceratopteris thalictroides)". May 2021.
  10. "Ceratopteris thalictroides - Useful Tropical Plants".

Further reading