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Biol Rev Camb Philos Soc
2022 Feb 01;971:273-298. doi: 10.1111/brv.12799.
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Sophistication in a seemingly simple creature: a review of wild holothurian nutrition in marine ecosystems.
Pierrat J
,
Bédier A
,
Eeckhaut I
,
Magalon H
,
Frouin P
.
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Holothurians are marine invertebrates that are among the most widespread benthic megafauna communities by both biomass and abundance in shallow-water and deep-sea ecosystems, their functions supporting important ecological services worldwide. Despite their simple appearance as sea cucumbers, holothurians show a wide range of feeding practices. However, information on what and how these animals eat is scattered and potentially confusing. We provide a comprehensive review of holothurian nutrition in coastal and deep-sea ecosystems. First, we describe morphological aspects of holothurian feeding and the ultrastructure of tentacles. We discuss the two processes for food capture, concluding that mucus adhesion is likely the main method; two mucous cells, type-1 and type-2, possibly allow the adhesion and de-adhesion, respectively, of food particles. Secondly, this review aims to clarify behavioural aspects of holothurian suspension- and deposit-feeding. We discuss the daily feeding cycle, and selective feeding strategies. We conclude that there is selectivity for fine and organically rich particles, and that feeding through the cloaca is also a route for nutrient absorption. Third, we provide a wide description of the diet of holothurians, which can be split into two categories: living and non-living material. We suggest that Synallactida, Molpadida, Persiculida, Holothuriida and Elasipodida, ingest the same fractions, and emphasise the importance of bacteria in the diet of holothurians.
Fig 1. Cumulative number of articles on “sea cucumber” for general publications and “bêche‐de‐mer” for those focusing on fisheries and aquaculture since 1950. Note the different scales for the two axes.
Fig 3. Fine ultrastructure of the holothurian tentacle. Digitate tentacle of (A) Thyonidium sp. and (B) Holothuria forskali. D, discs; P, Papillae; S, shaft. (C) Profile view of a five‐bud papilla. B, buds. (D) Arrangement of cilia and microvilli on bud surface. C, cilium; Mi, microvilli. (E, F) Diagrammatic sections through a whole tentacle (E) and through a two‐bud papilla (F). AL, ambulacral lumen; Co, connective tissue layer; EN, epineural nerve plexus; Ep, epidermis; HN, hyponeural nerve plexus; Me, mesothelium; Mu, muscles. Modified from Bouland et al. (1982) and McKenzie (1987).
Fig 4. Digestive tract morphology of (A) Holothuria atra and (B) H. tubulosa. BR, brown region of the anterior foregut; Cl, cloaca; CR, calcareous ring; DHP, dorsal haemal plexus; Fg, foregut; G, gonads; Hg, hindgut; LT, longitudinal tube of rete mirabile; MC, main channel of dorsal haemal plexus; Mg, midgut; RM, rete mirabile; RS, region of severance; RT, respiratory tree; S1;, sphincter 1; S₂, sphincter 2; S₃, sphincter 3; T, tentacles; TS, transverse sinuses; TV, tentacle vesicles; VP, vesicle of Poli; VS, ventral sinus. Modified from Trefz (1958) and Massin (1978, inspired by Cuénot, 1948).
Fig 5. Synthesis of food sources for the seven orders of holothurians. Order colours represent different trophic modes: blue, suspension‐feeders (Dendrochirotida); black, deposit‐feeders. The number of species per taxon is shown below the order name. The width of arrows corresponds to the proportion of food ingested per source. Colours of food sources vary with food type: green, living fraction; grey, non‐living fraction. Tentacle type for each taxon is represented by the drawings (see Fig. 2): digitate (unbranched); peltate (slightly branched); pinnate (highly branched); dendritic (ultra‐branched).
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