Capitella teleta

Last updated

Capitella teleta
Thumbnail Capitella teleta.jpg
Adult male
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Annelida
Class: Polychaeta
Family: Capitellidae
Genus: Capitella
C. teleta
Binomial name
Capitella teleta
Blake, Grassle & Eckelbarger, 2009

Capitella teleta is a small, cosmopolitan, segmented annelid worm. It is a well-studied invertebrate, which has been cultured for use in laboratories for over 30 years. [1] C. teleta is the first marine polychaete to have its genome sequenced. [2] [3]



Initial discovery

For many years researchers believed that Capitella capitata was the only representative of this genus that survived, and flourished, in polluted environments. After the oil spill that occurred near Cape Cod in West Falmouth, Massachusetts in 1969, researchers collected sediment and found an abundance of what they believed to be C. capitata. [4] [5] However, subsequent research showed that while the individuals collected from that region had very similar gross morphology, their life histories, methods of reproduction and genetics indicated there were at least six distinct species. Capitella species I, eventually described as Capitella teleta in 2009, was one of the initial species identified from these surveys. [6]


After 30 years of research on the group, Capitella teleta was officially described in 2009 by Blake et al. The species name is derived from the Greek word teleta, meaning "initiation". This word symbolizes that it was the first alternative Capitella species that was identified. [4]


A 2018 molecular phylogeny of the family Capitellidae established clear monophyly and showed 8 genera. [7] The phylogeny utilized 36 capitellid species and  combined data from 18S, 28S, H3, and COI gene sequences. This study also established Capitellidae as the sister group to Echiura. While the study attempted to map morphological characters to the molecular phylogeny, this was not phylogenetically informative and a more detailed re-evaluation of morphology could help to elucidate character trait evolution.

Taxonomic morphology

Capitella teleta has a narrow, segmented body with reduced parapodia and is red in color. There are nine anterior thoracic segments and many more abdominal segments. New segments are added throughout the lifespan from a posterior subterminal growth zone, called the posterior growth zone. Like other polychaetes, C. teleta has fine bristles or setae. Setae are segmentally repeated along the body, with morphologically distinct setae in the thoracic (hooded hooks) and abdominal segments (capillary setae). [4] This animal exhibits sexual dimorphism and males have dorsally-positioned genital spines on setigers 8-9 while females have paired ovaries in the abdominal segments. [8] Generally, there are separate sexes, however, hermaphroditism is possible when there are low densities of females. Males, females and hermaphrodites are of similar size (max size collected was a male that is 24 mm in length). [4] [9]



Capitella teleta lives in the shallow-water or intertidal marine environment. It is also found in salt marshes and is often found in high concentrations in disturbed soft sediments. It is a member of the infaunal benthic community. C. teleta burrows through the sediment by peristalsis, using its hydrostatic skeleton and contraction of longitudinal and circular muscles in the body wall. The thoracic segments of C. teleta also contain helical muscles that are proposed to generate additional force for burrowing. [10] Capitellids are commonly thought of as opportunistic in nature, due to their ability to inhabit and flourish in organically enriched marine sediments. [4] [5]

This organism is commonly found in sediments along the east and west coasts of North America. Additional reports have placed this group in the Mediterranean region as well as Japan. [4] [9]

Brood tube Thumbnail broodtube.jpg
Brood tube

Life history

Capitella teleta embryos and early larval stages develop in a brood tube that surrounds the mother. [11] The embryos are approximately 200 µm in diameter. [6] Over the course of approximately a week, the embryos develop into non-feeding larvae which form musculature, a centralized nervous system, two circular ciliary bands, two eye spots, segments, and setae. The larvae are non-feeding and the digestive system develops at a later stage than other organs.  Pre-metamorphosis larvae can be categorized into nine stages, with each stage lasting approximately one day. [12]  Upon further body elongation and gut maturation, the larvae emerge from the brood tube, and swim forward with a rotational turn via the beating of cilia organized within two circular bands, the prototroch and telotroch. [11] Larvae exhibit positive phototactic behavior in which they swim towards light, potentially an adaptation to aid in larval dispersal [13] [14] [15] C. teleta is an indirect developer and undergoes metamorphosis from a swimming larva into a burrowing juvenile.  Metamorphosis is characterized by cilia loss, body elongation, and crawling behavior. [16] Marine sediment functions as a cue to initiate metamorphosis into juvenile worms that thereafter grow into mature adults. [15] Competent larvae can be induced to metamorphose into juveniles when exposed to the B vitamins Nicotinamide (B3) and Riboflavin (B2), suggesting that these chemical compounds may be responsible for the inductive role of the marine sediment in larval metamorphosis. [17] The number of offspring in each brood tube can vary between 50 - 400 individuals, [6] and is influenced by food quality. [18]

After metamorphosis, the juveniles begin burrowing and feeding. The juvenile worms continue to grow and add segments during the eight weeks it takes to become sexually mature adults. Males and females can reproduce multiple times during their lifetime. Adults live approximately 12–14 weeks after maturation.

Stages of Capitella teleta larval development (Seaver et al., 2005) Thumbnail Cteleta stagingchart.jpg
Stages of Capitella teleta larval development (Seaver et al., 2005)


Capitella teleta feeds on the enriched sediment in which it burrows. C. teleta has a complex, regionalized alimentary canal consisting of a foregut, midgut and hindgut. [19] It ingests the sediment by everting its proboscis, which contains a ciliated, muscular dorsal pharynx. [20] Presence of a dorsal pharynx is uncommon in marine polychaetes, and this adaptation may have evolved independently in the family Capitellidae through selective pressures on feeding mode in the benthic marine niche they occupy. [20]


Stage 9 larvae Thumbnail Capitellalarva.jpg
Stage 9 larvae

A wide range of techniques have been developed to investigate C. teleta developmental processes. In 2006, the first study using whole mount in situ hybridization was published. [20] [21] This technique allows investigation of the expression and localization of specific mRNAs within a fixed sample. Immunohistochemistry was later developed as a way to visualize specific cell types in fixed specimens. [22] A microinjection protocol for uncleaved embryos and early cleavage stages was developed in 2010 and was used in a fate mapping study [23] to investigate the ultimate fate of blastomeres. [24] [25] Other useful techniques for studying early development of the embryo are targeted deletion of single cells with an infrared laser and blastomere isolation experiments. [11] [26] [13] [27] [28] Laser deletion was also utilized for the deletion of larval eyes at a later stage in development. [15] The development of microinjection techniques allowed for introduction of different nucleic acid constructs that can be injected into an uncleaved zygote. This includes use of gene perturbation techniques such as Morpholino knockdown and CRISPR-Cas9 mutagenesis, and methods for living imaging such as mRNA injection. [29] [30] [31] The development of each technique opens doors for new avenues of inquiry and experimentation and expands the number and complexity of questions C. teleta researchers can thoroughly investigate.


Like many species within Spiralia, C. teleta embryogenesis follows an unequal spiral cleavage program where blastomeres are born according to a predictable order, size and position. This shared stereotypic cleavage program allows for the identification of individual cells and there is a standard cell-nomenclature system. Additionally, individual cells can be microinjected with fluorescent dyes and their descendants tracked to determine the lineage of particular tissues and larval structures. Through this method, a comprehensive fate map was created for C. teleta. [23] In general, there is substantial similarity of cell fates between C. teleta and other Spiralia. [32] [23] [31] For instance, in C. teleta and several other spiralians, cells derived from the A, B, C, and D embryo quadrants respectively give rise to the left, ventral, right, and dorsal portions of the larval body. [23] However, the origin of mesoderm differs across species. In C. teleta, mesoderm is generated from cells called 3c and 3d that are derived from both the C and D embryo quadrants, but in the annelid Platynereis dumerilii and in several mollusks, trunk mesoderm is generated from a single cell 4d. [33] [34] [35] [36] [37]

The establishment of the dorsal-ventral axis during early embryological development has also been extensively studied in C. teleta. It is reported that micromere 2d, a cell that is born when the embryo has 16 cells, has organizing activity which enables it to induce dorsal-ventral polarity within the embryo. [13] Fate map studies have demonstrated that cell 2d gives rise to ectoderm in the larval trunk and pygidium in C. teleta, [24] while descendants of the first quartet micromeres give rise to structures in the larval head. When micromere 2d is laser ablated, 2d derived structures as well as dorsal-ventral organization in the head is lost. [13] This suggests a requirement for 2d to be present in order to induce the proper formation of the head along a dorsal-ventral axis. When micromeres 2d1 and 2d2, the immediate descendants of 2d, are both deleted, the resulting larvae retain dorsal-ventral organization within the head. [13] It was therefore concluded that in C. teleta micromere 2d has organizing activity in patterning the dorsal-ventral body axis. Furthermore, perturbation studies have shown that the dorsal-ventral axis is primarily patterned via the Activin/Nodal pathway. [38]


Many annelids possess the capability to regenerate their anterior, posterior, or both ends of their body. [39] C. teleta is capable of posterior regeneration. [40] [41] [42] Both juveniles and adults can regenerate their posterior halves quite well. A staging system has been established, describing the sequential regeneration events in juveniles of C. teleta. [43] The first stage of regeneration encompasses the first 24 hours following amputation or injury. This stage is marked by wound healing and a change in cell proliferation patterns. Wound healing occurs within 4–6 hours of amputation, as the circular muscles in the body wall contract, bringing the epithelium together to cover the wound. During this time, cell proliferation patterns are different from uncut animals; while cell proliferation is still observed throughout the body, there is a marked reduction at the wound site. In stage II, approximately 2 days after amputation, a small blastema forms that contains proliferating cells, and there is a diffuse network of neurites extending from the old ventral nerve cord tissue into the blastema. In stage III, approximately 3 days after amputation, the blastema becomes more organized as proliferating cells pack closely together in the newly formed tissue and multiple neurites condense into nerves. In stage IV, 5 days after amputation, there continues to be an increase in cell proliferation, but less so in the new tissue. The neural projections into the blastema become even more organized and patterned. Additionally, the posterior growth zone, pygidium, and hindgut reform. Finally, Stage 5 is marked by the presence and continued addition of new segments with differentiated tissues and ganglia. [44] [45] The entire regeneration process in C. teleta adults is completed within about two weeks [46] The rate of regeneration can vary among individuals, especially pertaining to health and nutrition intake.

Hox genes, patterning genes that regulate segment identity during development in many animals, and are expressed in the blastema of C. teleta during posterior regeneration. This suggests a role in the regeneration process, but the exact expression patterns do not make an obvious link to establishment of segment identity in newly formed tissue during regeneration. [42] The shift in Hox gene expression in the blastema during posterior regeneration is indicative of limited morphallaxis, in addition to epimorphic regeneration [42]

The regeneration of the germline in embryos has also been investigated. In early stage embryos, the germline precursor (cell 3D) was deleted using an infrared laser. 13% of screened larvae showed presence of multipotent progenitor cells (MPCs), indicating some regeneration of the germline. Furthermore, all juveniles two weeks post-metamorphosis have MPCs. Finally, almost all adult worms raised from treated embryos developed functional reproductive systems and produced offspring that developed into swimming larvae. [28]


Capitella teleta is an indicator species for environments contaminated with organic pollution. C. teleta has the ability to colonize these habitats rapidly with high growth rates. [4] [47] These characteristics have led to their use in various toxicological studies. Their population and/or individual- level responses to pollutant exposures have been investigated in various toxicants such as synthetic musk, [48] acetyl cedrene, [49] fluoranthene, [50] benzo[a]pyrene, [51] fluoxetine, [52] cadmium, [53] copper oxide nanoparticles, [54] and silver nanoparticles. [52] Recently, the effects of the fluoranthene-spiked sediments on the gut microbiome were investigated and several taxa of bacteria were identified; these taxa may play a role in the metabolism of fluoranthene. [55]


The genome of Capitella teleta was sequenced in concert with the owl limpet, Lottia gigantean, and the freshwater leech, Helobdella robusta , by the Joint Genome Institute in 2013. [2] [3] This was the first attempt at sequencing a marine polychaete and the sequencing and study of these three spiralian genomes provided an important perspective of early bilaterian evolutionary processes. Additionally, this work showed strong support for the monophyletic grouping of Lophotrochozoa.

The researchers found that when compared to other animal genomes, all three organisms possessed genome organization, gene structure and functional content that was more closely related to invertebrate deuterostome genomes than those of fellow invertebrate protostomes. C. teleta possesses a highly conserved and slowly evolving genome with respect to other metazoans. [3] [56]

Karyotype analysis revealed that C. teleta has 10 pairs of chromosomes. [57]

Related Research Articles

Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of stem cells in the adult organism.

Sipuncula Phylum of invertebrates, peanut worms

The Sipuncula or Sipunculida is a group containing about 162 species of bilaterally symmetrical, unsegmented marine worms. The name Sipuncula is from the genus name Sipunculus, and comes from the Latin siphunculus meaning a "small tube". Sipuncula seems to be closely related to Myzostomida, and Annelida.

Hemichordate Phylum of deuterostome animals

Hemichordata is a phylum of marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and include two main classes: Enteropneusta, and Pterobranchia. A third class, Planctosphaeroidea, is known only from the larva of a single species, Planctosphaera pelagica. The extinct class Graptolithina is closely related to the pterobranchs.

Segmentation in biology is the division of some animal and plant body plans into a series of repetitive segments. This article focuses on the segmentation of animal body plans, specifically using the examples of the taxa Arthropoda, Chordata, and Annelida. These three groups form segments by using a "growth zone" to direct and define the segments. While all three have a generally segmented body plan and use a growth zone, they use different mechanisms for generating this patterning. Even within these groups, different organisms have different mechanisms for segmenting the body. Segmentation of the body plan is important for allowing free movement and development of certain body parts. It also allows for regeneration in specific individuals.

Regeneration (biology) Biological process of renewal, restoration, and tissue growth

In biology, regeneration is the process of renewal, restoration, and tissue growth that makes genomes, cells, organisms, and ecosystems resilient to natural fluctuations or events that cause disturbance or damage. Every species is capable of regeneration, from bacteria to humans. Regeneration can either be complete where the new tissue is the same as the lost tissue, or incomplete where after the necrotic tissue comes fibrosis.

Somite Each of several blocks of mesoderm that flank the neural tube on either side in embryogenesis

The somites are a set of bilaterally paired blocks of paraxial mesoderm that form in the embryonic stage of somitogenesis, along the head-to-tail axis in segmented animals. In vertebrates, somites subdivide into the sclerotomes, myotomes, syndetomes and dermatomes that give rise to the vertebrae of the vertebral column, rib cage and part of the occipital bone; skeletal muscle, cartilage, tendons, and skin.


Somitogenesis is the process by which somites form. Somites are bilaterally paired blocks of paraxial mesoderm that form along the anterior-posterior axis of the developing embryo in segmented animals. In vertebrates, somites give rise to skeletal muscle, cartilage, tendons, endothelium, and dermis.

In developmental biology, cleavage is the division of cells in the early embryo. The process follows fertilization, with the transfer being triggered by the activation of a cyclin-dependent kinase complex. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size as the original zygote. The different cells derived from cleavage are called blastomeres and form a compact mass called the morula. Cleavage ends with the formation of the blastula.

Hox genes, a subset of homeobox genes, are a group of related genes that specify regions of the body plan of an embryo along the head-tail axis of animals. Hox proteins encode and specify the characteristics of 'position', ensuring that the correct structures form in the correct places of the body. For example, Hox genes in insects specify which appendages form on a segment, and Hox genes in vertebrates specify the types and shape of vertebrae that will form. In segmented animals, Hox proteins thus confer segmental or positional identity, but do not form the actual segments themselves.

Clitellata Class of annelid worms

The Clitellata are a class of annelid worms, characterized by having a clitellum - the 'collar' that forms a reproductive cocoon during part of their life cycles. The clitellates comprise around 8,000 species. Unlike the class of Polychaeta, they do not have parapodia and their heads are less developed.

Zona limitans intrathalamica

The zona limitans intrathalamica (ZLI) is a lineage-restriction compartment and primary developmental boundary in the vertebrate forebrain that serves as a signaling center and a restrictive border between the thalamus and the prethalamus.

Chaetopteridae Family of annelid worms

The Chaetopteridae are a family of marine filter-feeding polychaete worms that live in vertical or U-shaped tubes in tunnels buried in the sedimentary or hard substrate of marine environments. The worms are highly adapted to the hard tube they secrete. Inside the tube the animal is segmented and regionally specialized, with highly modified appendages on different segments for cutting the tunnel, feeding, or creating suction for the flow of water through the tube home. The modified segments for feeding are on the 12th segment from the head for members of this family.

engrailed is a homeodomain transcription factor involved in many aspects of multicellular development. First known for its role in arthropod embryological development, working in consort with the Hox genes, engrailed has been found to be important in other areas of development. It has been identified in many bilaterians, including the arthropods, vertebrates, echinoderms, molluscs, nematodes, brachiopods, and polychaetes. It acts as a "selector" gene, conferring a specific identity to defined areas of the body, and co-ordinating the expression of downstream genes.

Vasa is an RNA binding protein with an ATP-dependent RNA helicase that is a member of the DEAD box family of proteins. The vasa gene, is essential for germ cell development and was first identified in Drosophila melanogaster, but has since been found to be conserved in a variety of vertebrates and invertebrates including humans. The Vasa protein is found primarily in germ cells in embryos and adults, where it is involved in germ cell determination and function, as well as in multipotent stem cells, where its exact function is unknown.

Epimorphosis is defined as the regeneration of a specific part of an organism in a way that involves extensive cell proliferation of somatic stem cells, dedifferentiation, and reformation, as well as blastema formation. Epimorphosis can be considered a simple model for development, though it only occurs in tissues surrounding the site of injury rather than occurring system-wide. Epimorphosis restores the anatomy of the organism and the original polarity that existed before the destruction of the tissue and/or a structure of the organism. Epimorphosis regeneration can be observed in both vertebrates and invertebrates such as the common examples: salamanders, annelidas, and planarians.

Annelid Phylum of segmented worms

The annelids, also known as the ringed worms or segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecologies – some in marine environments as distinct as tidal zones and hydrothermal vents, others in fresh water, and yet others in moist terrestrial environments.

Evx1 is a mammalian gene located downstream of the HoxA cluster, which encodes for a homeobox transcription factor. Evx1 is a homolog of even-skipped (eve), which is a pair-rule gene that regulates body segmentation in Drosophila. The expression of Evx1 is developmentally regulated, displaying a biphasic expression pattern with peak expression in the primitive streak during gastrulation and in interneurons during neural development. Evx1 has been shown to regulate anterior-posterior patterning during gastrulation by acting as a downstream effector of the Wnt and BMP signalling pathways. It is also a critical regulator of interneuron identity.

Leech embryogenesis is the process by which the embryo of the leech forms and develops. The embryonic development of the larva occurs as a series of stages. During stage 1, the first cleavage occurs, which gives rise to an AB and a CD blastomere, and is in the interphase of this cell division when a yolk-free cytoplasm called teloplasm is formed. The teloplasm is known to be a determinant for the specification of the D cell fate. In stage 3, during the second cleavage, an unequal division occurs in the CD blastomere. As a consequence, it creates a large D cell on the left and a smaller C cell to the right. This unequal division process is dependent on actomyosin, and by the end of stage 3 the AB cell divides. On stage 4 of development, the micromeres and teloblast stem cells are formed and subsequently, the D quadrant divides to form the DM and the DNOPQ teloblast precursor cells. By the end stage 6, the zygote contains a set of 25 micromeres, 3 macromeres and 10 teloblasts derived from the D quadrant.

The protocerebrum is the first segment of the panarthropod brain.

Judith Grassle is a Professor Emeritus in the Department of Marine and Coastal Sciences at Rutgers University. Grassle is a benthic ecologist known for research on invertebrates, especially polychaete worms including the now-named Capitella teleta. Grassle became a Fellow of the American Association for the Advancement of Science in 1993.


  1. Blake, James A.; Grassle, Judith P.; Eckelbarger, Kevin J. (2009). "Capitella teleta, a new species designation for the opportunistic and experimental Capitella sp. I, with a review of the literature for confirmed records". Zoosymposia. 2: 25–53. doi: 10.11646/zoosymposia.2.1.6 .
  2. 1 2 "Home - Capitella sp. I ESC-2004". Retrieved 2017-04-21.
  3. 1 2 3 Simakov, Oleg; Marletaz, Ferdinand; Cho, Sung-Jin; Edsinger-Gonzales, Eric; Havlak, Paul; Hellsten, Uffe; Kuo, Dian-Han; Larsson, Tomas; Lv, Jie (2013-01-24). "Insights into bilaterian evolution from three spiralian genomes". Nature. 493 (7433): 526–531. Bibcode:2013Natur.493..526S. doi:10.1038/nature11696. ISSN   0028-0836. PMC   4085046 . PMID   23254933.
  4. 1 2 3 4 5 6 7 Grassle, JF; Grassle, JP (1974). "Opportunistic life histories and genetic systems in marine benthic polychaetes". Journal of Marine Research. 32: 253–284.
  5. 1 2 L., Sanders, Howard; Frederick, Grassle, J.; R., Hampson, George; S., Morse, Linda; Susan, Garner-Price; C., Jones, Carol (1980-05-01). "Anatomy of an oil spill : long-term effects from the grounding of the barge Florida off West Falmouth, Massachusetts". hdl:1912/3474.Cite journal requires |journal= (help)
  6. 1 2 3 Grassle, Judith P.; Grassle, J. Frederick (1976-01-01). "Sibling Species in the Marine Pollution Indicator Capitella (Polychaeta)". Science. 192 (4239): 567–569. Bibcode:1976Sci...192..567G. doi:10.1126/science.1257794. JSTOR   1741571. PMID   1257794.
  7. Tomioka, Shinri; Kakui, Keiichi; Kajihara, Hiroshi (October 2018). "Molecular Phylogeny of the Family Capitellidae (Annelida)". Zoological Science. 35 (5): 436–445. doi:10.2108/zs180009. hdl: 2115/75605 . ISSN   0289-0003. PMID   30298787.
  8. Eckelbarger KJ, Grassle JP: Ultrastructural differences in the eggs and ovarian follicle cells of the Capitella (Polychaeta) sibling species. Biol Bull 1983, 165:379-393.
  9. 1 2 Tomioka, Shinri; Kondoh, Tomohiko; Sato-Okoshi, Waka; Ito, Katsutoshi; Kakui, Keiichi; Kajihara, Hiroshi (2016-10-01). "Cosmopolitan or Cryptic Species? A Case Study of Capitella teleta (Annelida: Capitellidae)". Zoological Science. 33 (5): 545–554. doi:10.2108/zs160059. hdl: 2115/68333 . ISSN   0289-0003. PMID   27715419.
  10. Grill, S.; Dorgan, K. M. (2015-05-15). "Burrowing by small polychaetes - mechanics, behavior and muscle structure of Capitella sp". Journal of Experimental Biology. 218 (10): 1527–1537. doi: 10.1242/jeb.113183 . ISSN   0022-0949. PMID   25827841.
  11. 1 2 3 Pernet, Bruno; Amiel, Aldine; Seaver, Elaine C. (2012). "Effects of maternal investment on larvae and juveniles of the annelid Capitella teleta determined by experimental reduction of embryo energy content". Invertebrate Biology. 131 (2): 82–95. doi:10.1111/j.1744-7410.2012.00263.x. ISSN   1744-7410.
  12. Seaver, Elaine C.; Thamm, Katrin; Hill, Susan D. (July 2005). "Growth patterns during segmentation in the two polychaete annelids, Capitella sp. I and Hydroides elegans: comparisons at distinct life history stages". Evolution & Development. 7 (4): 312–326. doi:10.1111/j.1525-142X.2005.05037.x. ISSN   1520-541X. PMID   15982368.
  13. 1 2 3 4 5 Amiel, Aldine R.; Henry, Jonathan Q.; Seaver, Elaine C. (July 2013). "An organizing activity is required for head patterning and cell fate specification in the polychaete annelid Capitella teleta: New insights into cell–cell signaling in Lophotrochozoa". Developmental Biology. 379 (1): 107–122. doi: 10.1016/j.ydbio.2013.04.011 . PMID   23608454.
  14. Butman, CA; Grassle, JP; Buskey, EJ (1988). "Horizontal swimming and gravitational sinking of Capitella sp. I(Annelida: Polychaeta) larvae: implications for settlement". Ophelia. 29: 43–57.
  15. 1 2 3 Yamaguchi, Emi; Seaver, Elaine C. (December 2013). "The importance of larval eyes in the polychaete Capitella teleta : effects of larval eye deletion on formation of the adult eye". Invertebrate Biology. 132 (4): 352–367. doi: 10.1111/ivb.12034 .
  16. Biggers, William J.; Pires, Anthony; Pechenik, Jan A.; Johns, Eric; Patel, Priyam; Polson, Theresa; Polson, John (March 2012). "Inhibitors of nitric oxide synthase induce larval settlement and metamorphosis of the polychaete annelid Capitella teleta". Invertebrate Reproduction & Development. 56 (1): 1–13. doi:10.1080/07924259.2011.588006. ISSN   0792-4259.
  17. Burns, Robert T.; Pechenik, Jan A.; Biggers, William J.; Scavo, Gia; Lehman, Christopher (2014-11-12). Harder, Tilmann (ed.). "The B Vitamins Nicotinamide (B3) and Riboflavin (B2) Stimulate Metamorphosis in Larvae of the Deposit-Feeding Polychaete Capitella teleta: Implications for a Sensory Ligand-Gated Ion Channel". PLOS ONE. 9 (11): e109535. Bibcode:2014PLoSO...9j9535B. doi:10.1371/journal.pone.0109535. ISSN   1932-6203. PMC   4229104 . PMID   25390040.
  18. Marsh, A. G.; Gémare, A.; Tenore, K. R. (1989-09-01). "Effect of food type and ration on growth of juvenile Capitella sp. I (Annelida: Polychaeta): macro- and micronutrients". Marine Biology. 102 (4): 519–527. doi:10.1007/BF00438354. ISSN   1432-1793.
  19. Boyle, Michael J.; Seaver, Elaine C. (2008-01-03). "Developmental expression of foxA and gata genes during gut formation in the polychaete annelid, Capitella sp. I: Gut development in Capitella sp. I". Evolution & Development. 10 (1): 89–105. doi:10.1111/j.1525-142X.2007.00216.x. PMID   18184360.
  20. 1 2 3 Boyle, Michael J.; Seaver, Elaine C. (2009-08-31). "Evidence of a dorsal pharynx in the marine polychaete Capitella teleta (Polychaeta: Capitellidae)". Zoosymposia. 2 (1): 317–328. doi: 10.11646/zoosymposia.2.1.22 . ISSN   1178-9913.
  21. Seaver, Elaine C.; Kaneshige, Lori M. (2006-01-01). "Expression of 'segmentation' genes during larval and juvenile development in the polychaetes Capitella sp. I and H. elegans". Developmental Biology. 289 (1): 179–194. doi: 10.1016/j.ydbio.2005.10.025 . ISSN   0012-1606. PMID   16330020.
  22. Meyer, Néva P; Carrillo-Baltodano, Allan; Moore, Richard E; Seaver, Elaine C (December 2015). "Nervous system development in lecithotrophic larval and juvenile stages of the annelid Capitella teleta". Frontiers in Zoology. 12 (1): 15. doi:10.1186/s12983-015-0108-y. ISSN   1742-9994. PMC   4498530 . PMID   26167198.
  23. 1 2 3 4 Meyer, Néva P.; Boyle, Michael J.; Martindale, Mark Q.; Seaver, Elaine C. (2010-09-15). "A comprehensive fate map by intracellular injection of identified blastomeres in the marine polychaete Capitella teleta". EvoDevo. 1 (1): 8. doi:10.1186/2041-9139-1-8. ISSN   2041-9139. PMC   2949861 . PMID   20849573.
  24. 1 2 Meyer, Néva P.; Seaver, Elaine C. (2010-11-01). "Cell Lineage and Fate Map of the Primary Somatoblast of the Polychaete Annelid Capitella teleta". Integrative and Comparative Biology. 50 (5): 756–767. doi: 10.1093/icb/icq120 . ISSN   1540-7063. PMID   21558238.
  25. Meyer, Néva P.; Seaver, Elaine C. (November 2009). "Neurogenesis in an annelid: Characterization of brain neural precursors in the polychaete Capitella sp. I". Developmental Biology. 335 (1): 237–252. doi: 10.1016/j.ydbio.2009.06.017 . PMID   19540831.
  26. Carrillo-Baltodano, Allan M.; Meyer, Néva P. (2017-11-15). "Decoupling brain from nerve cord development in the annelid Capitella teleta: Insights into the evolution of nervous systems". Developmental Biology. 431 (2): 134–144. doi: 10.1016/j.ydbio.2017.09.022 . ISSN   0012-1606. PMID   28943340.
  27. Yamaguchi, Emi; Dannenberg, Leah C.; Amiel, Aldine R.; Seaver, Elaine C. (February 2016). "Regulative capacity for eye formation by first quartet micromeres of the polychaete Capitella teleta". Developmental Biology. 410 (1): 119–130. doi: 10.1016/j.ydbio.2015.12.009 . PMID   26702513.
  28. 1 2 Dannenberg, Leah C.; Seaver, Elaine C. (August 2018). "Regeneration of the germline in the annelid Capitella teleta". Developmental Biology. 440 (2): 74–87. doi: 10.1016/j.ydbio.2018.05.004 . PMID   29758179.
  29. Klann, Marleen; Seaver, Elaine C. (December 2019). "Functional role of pax6 during eye and nervous system development in the annelid Capitella teleta". Developmental Biology. 456 (1): 86–103. doi: 10.1016/j.ydbio.2019.08.011 . PMID   31445008.
  30. Neal, S.; de Jong, D. M.; Seaver, E. C. (2019-06-12). "CRISPR/CAS9 mutagenesis of a single r-opsin gene blocks phototaxis in a marine larva". Proceedings of the Royal Society B: Biological Sciences. 286 (1904): 20182491. doi:10.1098/rspb.2018.2491. ISSN   0962-8452. PMC   6571462 . PMID   31161907.
  31. 1 2 Seaver, Elaine C (2016-08-01). "Annelid models I: Capitella teleta". Current Opinion in Genetics & Development. Developmental mechanisms, patterning and evolution. 39: 35–41. doi:10.1016/j.gde.2016.05.025. ISSN   0959-437X. PMID   27318692.
  32. Lambert, J. David (2008). "Mesoderm in spiralians: the organizer and the 4d cell". Journal of Experimental Zoology Part B: Molecular and Developmental Evolution. 310B (1): 15–23. doi:10.1002/jez.b.21176. ISSN   1552-5015. PMID   17577229.
  33. Ackermann, Christian; Dorresteijn, Adriaan; Fischer, Albrecht (December 2005). "Clonal domains in postlarvalPlatynereis dumerilii (Annelida: Polychaeta)". Journal of Morphology. 266 (3): 258–280. doi:10.1002/jmor.10375. ISSN   0362-2525. PMID   16170805.
  34. Conklin, Edwin Grant (1897). "The embryology of crepidula, A contribution to the cell lineage and early development of some marine gasteropods". Journal of Morphology. 13 (1): 1–226. doi:10.1002/jmor.1050130102. hdl: 1912/605 . ISSN   1097-4687.
  35. Dictus, Wim J. A. G; Damen, Peter (1997-03-01). "Cell-lineage and clonal-contribution map of the trochophore larva of Patella vulgata (Mollusca)1Both authors contributed equally to this work.1". Mechanisms of Development. 62 (2): 213–226. doi: 10.1016/S0925-4773(97)00666-7 . ISSN   0925-4773. PMID   9152012.
  36. Lambert, J. David; Johnson, Adam B.; Hudson, Chelsea N.; Chan, Amanda (2016-08-08). "Dpp/BMP2-4 Mediates Signaling from the D-Quadrant Organizer in a Spiralian Embryo". Current Biology. 26 (15): 2003–2010. doi: 10.1016/j.cub.2016.05.059 . ISSN   0960-9822. PMID   27397892.
  37. Lambert, J. David; Nagy, Lisa M. (2001). "MAPK signaling by the D quadrant embryonic organizer of the mollusc Ilyanassa obsoleta". Development. 128 (1): 45–56. PMID   11092810.
  38. Lanza, Alexis R.; Seaver, Elaine C. (2018-03-01). "An organizing role for the TGF-β signaling pathway in axes formation of the annelid Capitella teleta". Developmental Biology. 435 (1): 26–40. doi: 10.1016/j.ydbio.2018.01.004 . ISSN   0012-1606. PMID   29337130.
  39. Bely, Alexandra E. (2006-08-01). "Distribution of segment regeneration ability in the Annelida". Integrative and Comparative Biology. 46 (4): 508–518. doi: 10.1093/icb/icj051 . ISSN   1540-7063. PMID   21672762.
  40. Hill, Susan D.; Savage, Robert M. (2009-01-01). Shain, Daniel H. (ed.). Annelids in Modern Biology . John Wiley & Sons, Inc. pp.  88–115. doi:10.1002/9780470455203.ch6. ISBN   9780470455203.
  41. Giani, Vincent C.; Yamaguchi, Emi; Boyle, Michael J.; Seaver, Elaine C. (2011-05-05). "Somatic and germline expression of piwi during development and regeneration in the marine polychaete annelid Capitella teleta". EvoDevo. 2: 10. doi:10.1186/2041-9139-2-10. ISSN   2041-9139. PMC   3113731 . PMID   21545709.
  42. 1 2 3 Jong, Danielle M. de; Seaver, Elaine C. (2016-02-19). "A Stable Thoracic Hox Code and Epimorphosis Characterize Posterior Regeneration in Capitella teleta". PLOS ONE. 11 (2): e0149724. Bibcode:2016PLoSO..1149724D. doi:10.1371/journal.pone.0149724. ISSN   1932-6203. PMC   4764619 . PMID   26894631.
  43. name=":22">Jong, Danielle M. de; Seaver, Elaine C. (2016-02-19). "A Stable Thoracic Hox Code and Epimorphosis Characterize Posterior Regeneration in Capitella teleta". PLOS ONE. 11 (2): e0149724. Bibcode:2016PLoSO..1149724D. doi:10.1371/journal.pone.0149724. ISSN   1932-6203. PMC   4764619 . PMID   26894631.
  44. de Jong, Danielle M.; Seaver, Elaine C. (2016-02-19). Schubert, Michael (ed.). "A Stable Thoracic Hox Code and Epimorphosis Characterize Posterior Regeneration in Capitella teleta". PLOS ONE. 11 (2): e0149724. Bibcode:2016PLoSO..1149724D. doi:10.1371/journal.pone.0149724. ISSN   1932-6203. PMC   4764619 . PMID   26894631.
  45. de Jong, Danielle M.; Seaver, Elaine C. (March 2018). "Investigation into the cellular origins of posterior regeneration in the annelid Capitella teleta". Regeneration. 5 (1): 61–77. doi:10.1002/reg2.94. PMC   5911572 . PMID   29721327.
  46. Bely, Alexandra E. (2006-08-01). "Distribution of segment regeneration ability in the Annelida". Integrative and Comparative Biology. 46 (4): 508–518. doi: 10.1093/icb/icj051 . ISSN   1540-7063. PMID   21672762.
  47. Grassle, Judith (1980), "Polychaete Sibling Species", in Brinkhurst, Ralph O.; Cook, David G. (eds.), Aquatic Oligochaete Biology, Springer US, pp. 25–32, doi:10.1007/978-1-4613-3048-6_3, ISBN   978-1-4613-3050-9
  48. Ramskov, Tina; Selck, Henriette; Salvitod, Daniel; Forbes, Valery E. (2009). "Individual- and population-level effects of the synthetic musk, hhcb, on the deposit-feeding polychaete, Capitella sp. I". Environmental Toxicology and Chemistry. 28 (12): 2695–2705. doi:10.1897/08-522.1. ISSN   1552-8618. PMID   19788341.
  49. Dai, Lina; Selck, Henriette; Salvito, Daniel; Forbes, Valery E. (2012). "Fate and effects of acetyl cedrene in sediments inhabited by different densities of the deposit feeder, Capitella teleta". Environmental Toxicology and Chemistry. 31 (11): 2639–2646. doi:10.1002/etc.1991. ISSN   1552-8618. PMID   22912158.
  50. Bach, Lis; Palmqvist, Annemette; Rasmussen, Lene Juel; Forbes, Valery E. (2005-09-30). "Differences in PAH tolerance between Capitella species: Underlying biochemical mechanisms". Aquatic Toxicology. 74 (4): 307–319. doi:10.1016/j.aquatox.2005.06.002. ISSN   0166-445X. PMID   16023227.
  51. Palmqvist, Annemette; Rasmussen, Lene Juel; Forbes, Valery E. (2008). "Relative impact of coexposure compared to single-substance exposure on the biotransformation and toxicity of benzo[a]pyrene and fluoranthene in the marine polychaete Capitella sp. I". Environmental Toxicology and Chemistry. 27 (2): 375–386. doi:10.1897/07-156R.1. ISSN   1552-8618. PMID   18348624.
  52. 1 2 Méndez, Nuria; Lacorte, Silvia; Barata, Carlos (2013-08-01). "Effects of the pharmaceutical fluoxetine in spiked-sediments on feeding activity and growth of the polychaete Capitella teleta". Marine Environmental Research. 89: 76–82. doi:10.1016/j.marenvres.2013.05.004. ISSN   0141-1136. PMID   23769338.
  53. Selck, Henriette; Decho, Alan W.; Forbes, Valery E. (1999). "Effects of chronic metal exposure and sediment organic matter on digestive absorption efficiency of cadmium by the deposit-feeding polychaete Capitella species I". Environmental Toxicology and Chemistry. 18 (6): 1289–1297. doi:10.1002/etc.5620180631. ISSN   1552-8618.
  54. Dai, Lina; Banta, Gary T.; Selck, Henriette; Forbes, Valery E. (2015-10-01). "Influence of copper oxide nanoparticle form and shape on toxicity and bioaccumulation in the deposit feeder, Capitella teleta". Marine Environmental Research. Particles in the Oceans: Implication for a safe marine environment. 111: 99–106. doi:10.1016/j.marenvres.2015.06.010. ISSN   0141-1136. PMID   26138270.
  55. Hochstein, Rebecca; Zhang, Qian; Sadowsky, Michael J.; Forbes, Valery E. (June 2019). "The deposit feeder Capitella teleta has a unique and relatively complex microbiome likely supporting its ability to degrade pollutants". Science of the Total Environment. 670: 547–554. Bibcode:2019ScTEn.670..547H. doi:10.1016/j.scitotenv.2019.03.255. PMID   30909032.
  56. Seaver, Elaine C (2016). "Annelid models I: Capitella teleta". Current Opinion in Genetics & Development. 39: 35–41. doi:10.1016/j.gde.2016.05.025. PMID   27318692.
  57. GRASSLE, J. P; GELFMAN, C. E.; MILLS, S. W. (1987). "Karyotypes of Capitella Sibling Species, and a Several Species in the Related Genera Capitellides and Capitomastus (Polychaeta)". Bulletin of the Biological Society of Washington (7): 77–88. ISSN   0097-0298.