Ceratopteris richardii

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Triangle water fern
C-Fern Culture Dish.jpg
"C-Fern" growing on agar media in cell culture dish
Status TNC G5.svg
Secure  (NatureServe)
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Family: Pteridaceae
Genus: Ceratopteris
Species:
C. richardii
Binomial name
Ceratopteris richardii

Ceratopteris richardii is a fern species belonging to the genus Ceratopteris , one of only two genera of the subfamily Parkerioideae of the family Pteridaceae. [1] 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 .

"C-Fern"

This particular species is of special scientific interest because a patented strain, called "C-Fern", was developed as a scientific aid and teaching tool in biology in 1995. [2] The use of "C-Fern" is facilitated by the fact that it grows readily in a cell-culture dish on agar media, reaching sexual maturity within 2–3 weeks of spore inoculation, with motile sperm cells being visible at this time. Over the course of about 6 weeks germination, sex determination and development of gametophytes, fertilization, embryogenesis, organogenesis, and sporophyte growth can all be observed, allowing an incredibly comprehensive study of the life cycle of homosporous ferns in a relatively short time period. [3] In addition, due to the small size of the plant many specimens can be observed growing simultaneously, allowing for larger sample sizes in research studies. Following the culture of "C-Fern" in dishes it can be transplanted to a dirt substrate, where it can be further allowed to grow and future generations can be used for subsequent studies.

Monilophytes are generally studied far less than other groups of plants and a full genome sequence is not yet available, however due to the development of "C-Fern" research into fern biology has been more prevalent and C. richardii has been used as a model organism to study vascular plant cell walls, alternation of generations (and associated mutations), genetics, population dynamics, and the effects of mitotic disrupter herbicides, among other topics. [4] [5] [6] Despite being genetically identical the inoculated spores can give rise to both hermaphrodites and male gametophytes, depending on the secretion of antheridiogen; this phenomenon has been used to study plant pheromones and the cascade of events that leads to epigenetic changes in ferns. [7] The ability to switch from bisexual to all male spores may provide an evolutionary advantage by promoting outbreeding. [8]

The use of C. richardii in genetic research studies has been valuable to understanding fern and plant evolution as a whole, and in 2019 "C-fern" became the first homosporous fern to have its genome partially assembled, thus acting as a reference genome to which other ferns can be compared. [9]

C. richardii spores germinated in space in 1999 on shuttle mission STS-93, making them one of the few plants to be grown in space. [10]

Development of "C-Fern" at 1 week after inoculation of spores onto agar media. Observed under bright field light microscopy. C-Fern Development - 1 Week.jpg
Development of "C-Fern" at 1 week after inoculation of spores onto agar media. Observed under bright field light microscopy.
Development of "C-Fern" at 3 weeks after inoculation of spores onto agar media. Observed under bright field light microscopy. C-Fern Development - 3 weeks.jpg
Development of "C-Fern" at 3 weeks after inoculation of spores onto agar media. Observed under bright field light microscopy.

Related Research Articles

Gametophyte 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">Flowering plant</span> Clade of seed plants that produce flowers

Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae, commonly called angiosperms. The term "angiosperm" is derived from the Greek words angeion and sperma ('seed'), and refers to those plants that produce their seeds enclosed within a fruit. They are by far the most diverse group of land plants with 64 orders, 416 families, approximately 13,000 known genera and 300,000 known species. Angiosperms were formerly called Magnoliophyta.

<span class="mw-page-title-main">Embryo</span> Multicellular diploid eukaryote in its earliest stage of development

An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm cell. The resulting fusion of these two cells produces a single-celled zygote that undergoes many cell divisions that produce cells known as blastomeres. The blastomeres are arranged as a solid ball that when reaching a certain size, called a morula, takes in fluid to create a cavity called a blastocoel. The structure is then termed a blastula, or a blastocyst in mammals.

Spore 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.

Fern Class of vascular plants

A fern is a member of a group of vascular plants that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except the lycopods, and differ from mosses and other bryophytes by being vascular, i.e., having specialized tissues that conduct water and nutrients and in having life cycles in which the branched sporophyte is the dominant phase. Ferns have complex leaves called megaphylls, that are more complex than the microphylls of clubmosses. Most ferns are leptosporangiate ferns. They produce coiled fiddleheads that uncoil and expand into fronds. The group includes about 10,560 known extant species. Ferns are defined here in the broad sense, being all of the Polypodiopsida, comprising both the leptosporangiate (Polypodiidae) and eusporangiate ferns, the latter group including horsetails, whisk ferns, marattioid ferns, and ophioglossoid ferns.

Alternation of generations Reproductive cycle of plants and algae

Alternation of generations is the predominant type of life cycle in plants and algae. It consists of a multicellular haploid sexual phase, the gametophyte, which has a single set of chromosomes alternating with a multicellular diploid asexual phase, the sporophyte which has two sets of chromosomes.

<i>Isoetes</i> Genus of vascular plants in the family Isoetaceae

Isoetes, commonly known as the quillworts, is the only extant genus of plants in the family Isoetaceae, which is in the class of lycopods. There are currently 192 recognized species, with a cosmopolitan distribution but with the individual species often scarce to rare. Some botanists split the genus, separating two South American species into the genus Stylites, although molecular data place these species among other species of Isoetes, so that Stylites does not warrant taxonomic recognition. Species of Isoetes virtually identical to modern forms have existed since the Jurassic epoch.

Germination Process by which an organism grows from a spore or seed

Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, ferns, bacteria, and the growth of the pollen tube from the pollen grain of a seed plant.

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

The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They 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'.

Embryophyte Subclade of green plants, also known as land plants

The Embryophyta, or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sister of the Zygnematophyceae/Mesotaeniaceae. The Embryophyta consist of the bryophytes plus the polysporangiophytes. Living embryophytes therefore include hornworts, liverworts, mosses, lycophytes, ferns, gymnosperms and flowering plants. The land plants have diplobiontic life cycles and is shown that the Charophycean green algae gave rise to land plants.

Green algae Paraphyletic group of autotrophic eukaryotes in the clade Archaeplastida

The green algae are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as sister of the Zygnematophyceae. Since the realization that the Embryophytes emerged within the green algae, some authors are starting to properly include them. The completed clade that includes both green algae and embryophytes is monophyletic and is referred to as the clade Viridiplantae and as the kingdom Plantae. The green algae include unicellular and colonial flagellates, most with two flagella per cell, as well as various colonial, coccoid and filamentous forms, and macroscopic, multicellular seaweeds. There are about 22,000 species of green algae. Many species live most of their lives as single cells, while other species form coenobia (colonies), long filaments, or highly differentiated macroscopic seaweeds.

Sporophyte 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. It develops from the zygote produced when a haploid egg cell is fertilized by a haploid sperm and each sporophyte cell therefore has a double set of chromosomes, one set from each parent. All land plants, and most multicellular algae, have life cycles in which a multicellular diploid sporophyte phase alternates with a multicellular haploid gametophyte phase. In the seed plants, the largest groups of which are the gymnosperms and flowering plants (angiosperms), the sporophyte phase is more prominent than the gametophyte, and is the familiar green plant with its roots, stem, leaves and cones or flowers. In flowering plants the gametophytes are very reduced in size, and are represented by the germinated pollen and the embryo sac.

Hornwort Division of non-vascular land plants with horn-shaped sporophytes

Hornworts are a group of non-vascular Embryophytes constituting the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of genetic information; the flattened, green plant body of a hornwort is the gametophyte stage of the plant.

Pteridophyte Paraphyletic group of spore-bearing vascular plants

A pteridophyte is a vascular plant that disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden. Ferns, horsetails, and lycophytes are all pteridophytes. However, they do not form a monophyletic group because ferns are more closely related to seed plants than to lycophytes. "Pteridophyta" is thus no longer a widely accepted taxon, but the term pteridophyte remains in common parlance, as do pteridology and pteridologist as a science and its practitioner, respectively. Ferns and lycophytes share a life cycle and are often collectively treated or studied, for example by the International Association of Pteridologists and the Pteridophyte Phylogeny Group.

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.

Leptosporangiate fern Subclass of ferns

The Polypodiidae, commonly called leptosporangiate ferns, formerly Leptosporangiatae, are one of four subclasses of ferns, and the largest of these, being the largest group of living ferns, including some 11,000 species worldwide. The group has also been treated as the class Pteridopsida or Polypodiopsida, although other classifications assign them a different rank. Older names for the group include Filicidae and Filicales, although at least the "water ferns" were then treated separately.

<i>Ceratopteris</i> Genus of aquatic plants

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.

Environmental 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.

Heterospory The production of spores of two different sizes and sexes by several groups of land plants

Heterospory is the production of spores of two different sizes and sexes by the sporophytes of land plants. The smaller of these, the microspore, is male and the larger megaspore is female. Heterospory evolved during the Devonian period from isospory independently in several plant groups: the clubmosses, the arborescent horsetails, and progymnosperms. This occurred as part of the process of evolution of the timing of sex differentiation.

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. C-fern official site
  3. Renzaglia, Karen Sue; Warne, Thomas R. (May 1995). "Ceratopteris: An Ideal Model System for Teaching Plant Biology". International Journal of Plant Sciences. 156 (3): 385–392. doi:10.1086/297260. ISSN   1058-5893. S2CID   85418664.
  4. Spiro, Mark D.; Knisely, Karin I. (March 2008). Sundberg, Marshall (ed.). "Alternation of Generations and Experimental Design: A Guided-Inquiry Lab Exploring the Nature of the her1 Developmental Mutant of Ceratopteris richardii (C-Fern)". CBE: Life Sciences Education. 7 (1): 82–88. doi: 10.1187/cbe.07-82-88 . ISSN   1931-7913. PMC   2262118 . PMID   18316811.
  5. Hoffman, J. C.; Vaughn, K. C. (March 1996). "Spline and flagellar microtubules are resistant to mitotic disrupter herbicides". Protoplasma. 192 (1–2): 57–69. doi:10.1007/BF01273245. ISSN   0033-183X. S2CID   19844471.
  6. Leroux, Olivier; Eeckhout, Sharon; Viane, Ronald L. L.; Popper, Zoë A. (2013). "Ceratopteris richardii (C-fern): a model for investigating adaptive modification of vascular plant cell walls". Frontiers in Plant Science. 4: 367. doi: 10.3389/fpls.2013.00367 . ISSN   1664-462X. PMC   3779834 . PMID   24065974.
  7. Atallah, Nadia M.; Vitek, Olga; Gaiti, Federico; Tanurdzic, Milos; Banks, Jo Ann (July 2018). "Sex Determination in Ceratopteris richardii Is Accompanied by Transcriptome Changes That Drive Epigenetic Reprogramming of the Young Gametophyte". G3: Genes, Genomes, Genetics. 8 (7): 2205–2214. doi:10.1534/g3.118.200292. ISSN   2160-1836. PMC   6027899 . PMID   29720393.
  8. Timothy Walker, Plants: A Very Short Introduction. Oxford University Press, 2012. p. 49
  9. Marchant, D. Blaine; Sessa, Emily B.; Wolf, Paul G.; Heo, Kweon; Barbazuk, W. Brad; Soltis, Pamela S.; Soltis, Douglas E. (December 2019). "The C-Fern (Ceratopteris richardii) genome: insights into plant genome evolution with the first partial homosporous fern genome assembly". Scientific Reports. 9 (1): 18181. Bibcode:2019NatSR...918181M. doi:10.1038/s41598-019-53968-8. ISSN   2045-2322. PMC   6890710 . PMID   31796775.
  10. Salmi, ML; Roux, SJ (2008). "Gene expression changes induced by space flight in single-cells of the fern Ceratopteris richardii". Planta. 229 (1): 151–9. doi:10.1007/s00425-008-0817-y. PMID   18807069. S2CID   30624362.