Terebratalia transversa

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Terebratalia transversa
Terebratalia transversa 141510036.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Brachiopoda
Class: Rhynchonellata
Order: Terebratulida
Family: Terebrataliidae
Genus: Terebratalia
Species:
T. transversa
Binomial name
Terebratalia transversa
(Sowerby, 1846)

Terebratalia transversa or the North Pacific Lampshell is a species of marine brachiopod in the family Terebrataliidae. [1] [2] A two-valved shelled species, they are most frequently found in tidal habitats in the Pacific Northwest of the United States. [3] [4] [5]

Contents

Terebratalia transversa radial symmetry development Terebratalia transversa cleavage.svg
Terebratalia transversa radial symmetry development

Life history

Larval stage

The species belongs to the deuterostome class, developing their embryos through radial cleavage. [4] Researchers find that the mesoderm found in Terebertalia larvae comes from the endomesoderm and develops close to the vegetal blastopore. [4]

Its larval features include the apical organ, apical ciliary tuft, and bands of ciliated sections. [6] The features of the apical organ suggest an evolutionary reconstruction of brains commonly seen in bilaterians. [6] During the development of this apical organ, the ciliated sections form together into a tuft with recessed rootlets that is made up of acetylated α-tubulin. [6]

The larvae are lecithotrophic and typically spend four days as juveniles until they undergo metamorphosis. In past studies, it appeared that the ideal temperature for T. transversa development is 11 °C. The first muscles to develop in the species are the pedicle and mantle muscles. Shortly after, they develop four setae pouches which later connect to their developing circular mantle. They are free-swimming and during larval stages, carry two cell types in their epithelium: lobate and vesicular cells. The lobate cells develop a thin layer of undescribed material while the vesicular cells create a sheet dense in electrons, that will eventually become the foundation of their developing periostracum. [7]

In late-stage larvae, the species has their pedicle muscles connected to their central mantle which then is connected to the setae pouches along with an apical musculature. At this point, the species is referred to as a three-lobed larva. [8] Late stage T. transversa larvae also develop pigmented ocelli that develop on the dorsal side of their apical lobe. [6]

Prior to the brain neuropile that develops in the adult stage of the species, the late larval stages contain a nervous system of T. transvesa consists of three neural domains, one that is located ventrally to the mantle lobe of the larvae and two that are located anteriorly. [6]

Juveniles develop quickly after their A-P axis emerges due to Hox gene expression in the event referred to as "metamorphosis." Two days after this metamorphosis, T. transversa display adult morphology. The point at which they are referred to as adults is when they have a clearly separated body from their old two-valved juvenile shell attached via a posterior pedicle. [9]

Adult stage

Terebratalia transversa adults can grow up to an average of 50 millimeters long. They have a distinctive, thin two-layer calcitic shell. Like most invertebrates, this species cannot survive in water temperatures higher than 35 °C. [3] During adulthood they are filter feeding animals rather than their non-feeding lifestyle as larvae. [9]

They contain a lophophore, a ciliated feeding organ that has a similar appearance to an external tentacle. They also have two-valved shells connected by a singular tooth through a socket hinge. [10]

Inside of a North Pacific Lampshell showing the socket hinge and intestine with striated muscle fibers Terebratalia transversa 115544343.jpg
Inside of a North Pacific Lampshell showing the socket hinge and intestine with striated muscle fibers

T. transversa has a nervous system that consists of a brain neuropile found dorsally to a slit found on the surface of their bodies. The musculature consists of rudiments of pedicle adjustors, shell diductors, and shell adductors. It is noted that in recently metamorphic juveniles, musculature can remodel variably. The most significant muscle morphology in the species is referred to as the mantle margin, the tissues responsible for the development of their adult shell. [9]

T. transversa has musculature that composes their intestine and has tentacles of which contain multiple striated muscle fibers. [8] They contain no anus nor an articulated valve hinge. [9]

The calcitic shell is an identifying aspect of the species, however, their shells vary extremely in appearance between individuals within a population. In developing their shells, the species develops additions of carbonate to their exterior and later, during adulthood, their shells begin secreting epithelium. [3] [9] Their shells are divided into two classifications, the primary layer and the secondary layer. The outermost part of the shell is the primary layer which contains transversal sections of fibers pointing in various directions. The secondary shell layer is typically thicker and contain similar transversal fibers. In the both layers, there are punctuations dotting the shell appearing as slits in the shell tissue. [3] During shell development, these fibers, which are also composed of calcite, can rotate against their orientation gradient which strengthens the overall shell structure. The composition and overall shape of shells are heavily dependent on seawater molecular composition and tidal habitats. [3]

Calcite shell showing distinct growing periods separated by striations Terebratalia transversa (brachiopod shell) (modern; offshore California, USA) 3.jpg
Calcite shell showing distinct growing periods separated by striations

Varying morphology

Presenting accurate morphological details of Terebratalia transversa has proven to be difficult as studies have shown that the species is highly variable in their morphology. The overall body form of the species can vary immensely with some appearing as prolate spheroids and others as oblate spheroids. [11] Another difficultly when examining traits of the species is the variance between shell features of the species, specifically the smooth and ribbed shells found within this species. It has been proposed that meristic traits are not applicable in the homoegenous classification of T. transversa. [11]

Habitat and distribution

Significant populations of Terebratalia transversa are located in the waters near the Pacific Northwest in the United States. Specifically, scientists have collected the species by dredging in the area around the San Juan Channel off the coast of Washington, USA. [4] The species resides in tidal and sub-tidal habitats. [5] The molecular composition of these habitats affect heavily the size of shells as the carbon-oxygen isotope ratios in the seawater surrounding them will affect their shell growth rates. [3]

Fossil and genetic analysis

Due to the very thin calcitic shell of Terebratalia, the primary layer of these shells very rarely fossilize. The shells of this species, when fossilized, help researchers find preserved isotopic signals that aid in uncovering molecular compositions of an old Paleozoic ocean. [3]

Somewhat recently, researchers at the University of Michigan completed the mitochondrial genome of Terebratalia transversa. They uncovered that the mtDNA encodes for two rRNAs, thirteen proteins, and twenty-two tRNAs. The genetic information used to synthesize these compounds is comprised in 37 genes in the mitochondria. [10] Something uncommon among other described mtDNAs in animals, is that the genes of T. transversa are transcribed from the same strand. [10]

Most of the known morphology of T. transversa comes from research based on confocal laser scanning microscopy (CLSM) in conjunction with both scanning and transmission electron microscopy.

Related Research Articles

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<span class="mw-page-title-main">Trematoda</span> Class of parasitic flatworms

Trematoda is a class of flatworms known as flukes or trematodes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive host, where the flukes sexually reproduce, is a vertebrate. Infection by trematodes can cause disease in all five traditional vertebrate classes: mammals, birds, amphibians, reptiles, and fish.

<span class="mw-page-title-main">Veliger</span> Larval stage of some snails

A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells.

<span class="mw-page-title-main">Lingulata</span> Class of marine lamp shells

Lingulata is a class of brachiopods, among the oldest of all brachiopods having existed since the Cambrian period. They are also among the most morphologically conservative of the brachiopods, having lasted from their earliest appearance to the present with very little change in shape. Shells of living specimens found today in the waters around Japan are almost identical to ancient Cambrian fossils.

<span class="mw-page-title-main">Craniata (brachiopod)</span> Class of marine lamp shells

Craniata is a class of brachiopods originating in the Cambrian period and still extant today. It is the only class within the subphylum Craniiformea, one of three major subphyla of brachiopods alongside linguliforms and rhynchonelliforms. Craniata is divided into three orders: the extinct Craniopsida and Trimerellida, and the living Craniida, which provides most information on their biology. Living members of the class have shells which are composed of calcite, though some extinct forms my have aragonite shells. The shells are inarticulate and are usually rounded in outline. There is no pedicle; the rear edge of the body cavity is a smooth and flat wall perforated by the anus. This class of brachiopods has an unsupported lophophore with only a single row of tentacles. In the absence of a pedicle, the shell is usually attached directly to a hard substrate. Many craniiforms are encrusting animals which attach directly to the shell of another animal, usually another brachiopod. The plicae from the host brachiopod will then appear within the shell of the craniiform.

The Obolellata are a class of Rhynchonelliform brachiopods with two orders, Obolellida and Naukatida. They are essentially restricted to the lower-middle Cambrian.

<i>Lingula</i> (brachiopod) Genus of brachiopods within the class Lingulata

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<span class="mw-page-title-main">Brachiopod</span> Phylum of marine animals also known as lamp shells

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<span class="mw-page-title-main">Acrotretida</span> Extinct order of brachiopods

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<span class="mw-page-title-main">Rhynchonelliformea</span> Subphylum of brachiopods

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References

  1. "WoRMS - World Register of Marine Species - Terebratalia transversa (Sowerby, 1846)". www.marinespecies.org. Retrieved 2023-03-25.
  2. taxonomy. "Taxonomy browser (Terebratalia transversa)". www.ncbi.nlm.nih.gov. Retrieved 2023-03-25.
  3. 1 2 3 4 5 6 7 Auclair, Anne-Cécile; Joachimski, Michael M; Lécuyer, Christophe (2003-12-15). "Deciphering kinetic, metabolic and environmental controls on stable isotope fractionations between seawater and the shell of Terebratalia transversa (Brachiopoda)". Chemical Geology. 202 (1): 59–78. Bibcode:2003ChGeo.202...59A. doi:10.1016/S0009-2541(03)00233-X. ISSN   0009-2541.
  4. 1 2 3 4 Passamaneck, Yale J.; Hejnol, Andreas; Martindale, Mark Q. (2015-04-11). "Mesodermal gene expression during the embryonic and larval development of the articulate brachiopod Terebratalia transversa". EvoDevo. 6 (1): 10. doi: 10.1186/s13227-015-0004-8 . ISSN   2041-9139. PMC   4404124 . PMID   25897375.
  5. 1 2 Hughes, W. W.; Rosenberg, G. D.; Tkachuck, R. D. (1988-07-01). "Growth increments in the shell of the living brachiopod Terebratalia transversa". Marine Biology. 98 (4): 511–518. doi:10.1007/BF00391542. ISSN   1432-1793. S2CID   85016150.
  6. 1 2 3 4 5 Santagata, Scott; Resh, Carlee; Hejnol, Andreas; Martindale, Mark Q.; Passamaneck, Yale J. (2012-01-24). "Development of the larval anterior neurogenic domains of Terebratalia transversa (Brachiopoda) provides insights into the diversification of larval apical organs and the spiralian nervous system". EvoDevo. 3 (1): 3. doi: 10.1186/2041-9139-3-3 . ISSN   2041-9139. PMC   3314550 . PMID   22273002.
  7. Stricker, Stephen A.; Reed, Christopher G. (1985). "The ontogeny of shell secretion inTerebratalia transversa (brachiopoda, articulata) I. Development of the mantle". Journal of Morphology. 183 (3): 233–250. doi:10.1002/jmor.1051830302. ISSN   0362-2525. PMID   4039009. S2CID   45063595.
  8. 1 2 Altenburger, Andreas; Wanninger, Andreas (2009-02-03). "Comparative larval myogenesis and adult myoanatomy of the rhynchonelliform (articulate) brachiopods Argyrotheca cordata, A. cistellula, and Terebratalia transversa". Frontiers in Zoology. 6 (1): 3. doi: 10.1186/1742-9994-6-3 . ISSN   1742-9994. PMC   2645390 . PMID   19192287.
  9. 1 2 3 4 5 Gąsiorowski, Ludwik; Hejnol, Andreas (2019-01-08). "Hox gene expression in postmetamorphic juveniles of the brachiopod Terebratalia transversa". EvoDevo. 10 (1): 1. doi: 10.1186/s13227-018-0114-1 . ISSN   2041-9139. PMC   6325747 . PMID   30637095.
  10. 1 2 3 Helfenbein, Kevin; Brown, Wesley; Boore, Jeffrey (September 2001). "The Complete Mitochondrial Genome of the Articulate Barchiopod Terebratalia transversa". academic.oup.com. Retrieved 2023-04-18.
  11. 1 2 Paine, Robert (1969). "Growth and Size Distribution of the Brachiopod Terebratalia transversa Sowerby". University of Hawai'i Press. 23 (3): 337–343.