A morphohistological and histochemical study of hatchery-reared European hake, Merluccius merluccius (Linnaeus, 1758), during the lecitho-exotrophic larval phase


  • Juan B. Ortiz-Delgado Instituto de Ciencias Marinas de Andalucía - CSIC
  • José Iglesias Instituto Español de Oceanografía
  • Francisco J. Sánchez Instituto Español de Oceanografía
  • Rosa Cal Instituto Español de Oceanografía
  • M. Jesus Lago Instituto Español de Oceanografía
  • Juan J. Otero Instituto Español de Oceanografía
  • Carmen Sarasquete Instituto de Ciencias Marinas de Andalucía - CSIC




European hake, Merluccius merluccius, larvae, histology, histochemistry, ontogeny, yolk sac, endo-exotrophic feeding, visual acuity


The larval development of reared European hake, Merluccius merluccius (Linnaeus, 1758), during the lecithotrophic phase, from hatching until 5 days post-hatching (dph), and throughout the endo-exotrophic feeding phase (6-10 dph) was studied by histology and histochemistry. Many crucial morphological, cellular and tissular changes were observed during both feeding phases, mostly those related to digestive and visual ontogenetic events, such as differentiation of buccopharyngeal cavity and eye development (at hatching); pigmentation and differentiation of cone-photoreceptors (4 dph); opening of the mouth and anus, appearance of intestinal valves (5-6 dph); presence of buccopharyngo-esophageal (5-6 dph) and intestinal (9 dph) mucous cells; folding of intestinal mucosa (6 dph); development of regional specific digestive musculature (6 dph); typical structure and functionality of the liver (sinusoids, biliary and pancreatic ducts, glycogen, protein and lipid reserves) (4-6 dph); and characteristic acinar distribution pattern of eosinophilic zymogen granules of the exocrine pancreas (6 dph). Between 9 and 10 dph, the hake larvae showed evident signs of lipid absorption within enterocytes of the anterior intestinal region and a remarkable process of pynocitosis and intracellular digestion was detected in the posterior intestine (supranuclear inclusions or acidophilic protein vesicles). In hake larvae at 10 dph, a proliferation of renal tubules, spleen differentiation and gill development, as well as the presence of the first thyroid follicle, were clearly distinguished. At this time, stomach gastric gland differentiation was not detected and endocrine pancreas and gill lamellae were not evidenced. However, and interestingly, swim bladder and eyes (developing rods) were well differentiated in larval development from 9 dph onwards. In summary, in European hake larval development during the endo-exogenous feeding phase and especially at 9 to 10 dph, most systems, organs and tissues were well differentiated. Particularly, digestive and visual systems were developed and physiologically functional some days before the start of the exogenous feeding phase.


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Alheit J., Picher T.J. (eds.) 1995. Hake: fisheries, ecology and markets. Fish and fisheries series 15. Chapman & Hall, London, 487 pp.

Arneri E., Morales-Nin B. 2000 . Aspects of the early history of the European hake from the central Adriatic. J. Fish Biol. 46: 1368-1380. http://dx.doi.org/10.1111/j.1095-8649.2000.tb02149.x

Bjelland R.M., Skiftesvik A.B. 2006. Larval development in European hake (Merluccius merluccius L) reared in a semi-intensive culture. Aq. Res. 37: 1117-1129. http://dx.doi.org/10.1111/j.1365-2109.2006.01536.x

Blaxter J.H.S. 1986. Development of sense organs and behaviour of teleost larvae with special reference to feeding and predator avoidance. Trans. Am. Fish. Soc. 115: 98-114. http://dx.doi.org/10.1577/1548-8659(1986)115<98:NLFCDO>2.0.CO;2

Bozzano A., Catalan I.A. 2002. Ontogenetic changes in the retinal topography of the European hake, Merluccius merluccius: implications for feeding and depth distribution. Mar. Biol. 141: 549-559. http://dx.doi.org/10.1007/s00227-002-0840-7

Branchet T., Bremiller R. 1984. The development of photoreceptors in the zebrafish, Brachiodanio rerio. J. Comp. Neur. 224: 116-122. http://dx.doi.org/10.1002/cne.902240110 PMid :6715575

Bustos C.A., Landaeta F., Bay-Schmith E., Lewis R., Moraga X. 2007. Effects of temperature and lipid droplet adherence on mortality of hatchery-reared southern hake, Merluccius merluccius larvae. Aquaculture 270: 535-540. http://dx.doi.org/10.1016/j.aquaculture.2007.03.032

Carvalho L., Hesberg C.P. 2010. The yolk syncytial layer in early zebrafish development. Trends Cell Biol. 20: 586-592. http://dx.doi.org/10.1016/j.tcb.2010.06.009 PMid :20674361

Casey J., Pereiro J. 1995. European hake (M. merluccius) in the Nort-east Atlantic. In. Alheit J and Pitcher, T.J (eds.), Hake, fisheries, ecology and markets. Fish and fisheries series 15. Chapman & Hall, London, pp: 142-147.

Deplano M., Diaz J.P., Connes R., Kentouri-Divamach M., Cavalier F. 1991. Appearance of lipid-absorption capacities in larvae of the sea bass, Dicentrarchus labrax during transition to the exotrophic phase. Mar. Biol. 108: 361-371. http://dx.doi.org/10.1007/BF01313645

Dinis M.T., Ribeiro L., Soares F., Sarasquete C. 1999. A review on the cultivation potential of Solea senegalensis in Spain and Portugal. Aquaculture 176: 27-38. http://dx.doi.org/10.1016/S0044-8486(99)00047-2

Falk-Petersen I.B. 2005. Comparative organ differentiation during early life stages of marine fish. Fish Sellfish Inmunol. 19: 397-412. http://dx.doi.org/10.1016/j.fsi.2005.03.006

FAO. 2005. Review of the state of world marine fishery resources. FAO. Fish Tech. Pap., 457: 235 pp.

Gisbert E., Ortiz-Delgado J.B., Sarasquete C. 2008. Nutritional cellular biomarkers in early life stages of fish. Histol. Histopathol. 23: 1525-1539. PMid :18830938

Groison, A.L. 2010. Male reproductive biology of the European hake, Merluccius merluccius. Ph. D. thesis, Univ. Bergen (Norway), 88 pp.

Haacke C., Martin H., Melzer R.R., Gebhart H., Smola U. 2001. Fine structure and development of the retina of the grenadier anchovy, Coilia nasus (Engraulidae, Cupleiformes). J. Morphol. 248: 41-55 http://dx.doi.org/10.1002/jmor.1019 PMid :11268057

Hadedorn M., Fernald R.D. 1992. Retinal growth and cell addition during embryogenesis of the teleost, Haplochromis burtoni. J. Comp. Neur. 321: 193-208. http://dx.doi.org/10.1002/cne.903210203 PMid :1380013

Iglesias J., Lago M.J., Sánchez F.J., Cal R. 2010. Short communication: Capture, transport and acclimation to captivity of the European hake, Merluccius merluccius: preliminary date on feeding and growth. Aq. Res. 41: 607-609. http://dx.doi.org/10.1111/j.1365-2109.2009.02365.x

Kawamura G. 1984. The visual cell morphology of Pagrus pagrus and its adaptative changes with shift from pelagic and benthic habits. Bull. Jpn. Soc. Sci. Fish. 50: 1975-1980. http://dx.doi.org/10.2331/suisan.50.1975

Kawamura G., Washiyama N. 1989. Ontogenetic changes in behaviour and sense organ morphogenesis in largemouth bass and Tilapia nilotica. Trans. Am. Fish. Soc. 118: 203-213. http://dx.doi.org/10.1577/1548-8659(1989)118<0203:OCIBAS>2.3.CO;2

Kawamura G., Mukay Y., Ohta H. 1984. Changes in the visual threshold with development of rods in ayu, Pecoglossus altivelis. Nip. S. Gak. 50: 2133.

Kawamura G., Masuma S., Tezuka N., Koiso M., Jimbo J., Namba K. 2003. Morphogenesis of the sense organs in the bluefin tuna, Thunnus orientalis. In: Browman and Skiftesvik (eds.), The big fish band. N-5817. Bergen, Norway, pp: 123-135.

Kuz’mina V.V., Gel’man G.A. 1998. Traits in development of the digestive functions in fishes. J. Ichthtyol. 38: 106-115.

Land M.F., Nilsson D.E. 2002. Animal eyes. Oxford University Press, Oxford, 244 pp.

Last J.M. 1980. The food of twenty species of fish larva in the west-central North Sea. Fish Res. 60: 1-44.

Lentz T.L., Trinkaus P. 1967. A fine structure of cytodifferentiation during cleavage, blastula and gastrula stages of Fundulus heteroclitus. J. Cell Biol. 32: 121-137. http://dx.doi.org/10.1083/jcb.32.1.121 PMid :10976205    PMCid:2107097

Lloris D., Matallanas J., Oliver P. 2005. Species catalogue for fishery Purposes. Hakes of the World. FAO, Roma, 57 pp.

Martoja, R. Martoja-Pierson M. 1970. Técnicas de Histología Animal. Ed Toray-Masson, Barcelona (Spain), 350 pp.

Margulies S. 1997. Development of the visual system and inferred performance capabilities of larval and early juvenile scombrids. Mar. Fresh. Behav. Physiol. 30: 203-213. http://dx.doi.org/10.1080/10236249709379018

Mas-Riera J. 1991. Changes during growth in the retinal structure of three hake species, Merluccius (Teleostei; Gadiformes) in relation to their depth distribution and feeding. J. Exp. Mar. Biol. Ecol. 152: 91-104. http://dx.doi.org/10.1016/0022-0981(91)90137-L

Morales-Nin B., Bjelland R.M., Moksness E. 2005. Otolith microstructure of a hatchery reared European hake (Merluccius meruccius). Fish. Res. 74: 300-305. http://dx.doi.org/10.1016/j.fishres.2005.03.001

Morote E., Olivar M.P., Pankhurst P.M., Villate F., Uriarte I. 2008. Tropic ecology of blue tuna Auxis rockei larvae and ontogeny of feeding-related organs. Mar. Ecol. Prog. Ser. 353: 243-254. http://dx.doi.org/10.3354/meps07206

Morote E., Olivar M.P., Bozzano A., Villate F., Uriarte I. 2011. Feeding selectivity in larvae of the European hake (Merluccius merluccius) in relation to ontogeny and visual capabilities. Mar. Biol. 158(6): 1349-1361. http://dx.doi.org/10.1007/s00227-011-1654-2

Murua H., Saborido-Rey F. 2003. Female reproductive strategies of marine fish species of the North Atlantic. J. Northw. Atl. Sish. Sci. 33: 23-31 http://dx.doi.org/10.2960/J.v33.a2

Ortiz-Delgado J.B., Ruane N.M., Pouçao P., Dinis M.T., Sarasquete C. 2006. Thyroid gland development in Senegales sole, Solea senegalensis Kaup 1858 during early life stages: a histochemical and immunohistochemical approach. Aquaculture 260: 346-353. http://dx.doi.org/10.1016/j.aquaculture.2006.06.006

Ortiz-Delgado J.B., Porcelloni S., Fossi C., Sarasquete C. 2008. Histochemical characterization of the swordfish, Xiphias gladius oocytes. Sci. Mar. 72(3): 549-568.

Olivar M.P., Quílez G., Emelianov M. 2003. Spatial and temporal distribution and abundance of European hake, Merluccius merluccius, eggs and larvae in the Catalan coast (NW Mediterranean). Fish Res. 60: 321-331. http://dx.doi.org/10.1016/S0165-7836(02)00139-X

Oliver P., Massutí E. 1995. Biology and fisheries of western Mediterranean hake (Merluccius merluccius). In: Alheit J, Pitcher TJ (eds.), Hake: fisheries, ecology and markets. Chapman & Hall, London, pp: 181-202.

Omura Y., Uematsu K., Tachiki H., Furukawa K., Satoh H. 1997. Cone cells appear also in retina of eel larvae. Fish. Sci. 63: 1052-1053.

Palomera I., Olivar M.P., Morales-Nin B. 2005. Larval development and growth of the European hake in the NW Mediterranean. Sci. Mar. 69: 221-258.

Pearse, A.G.E. 1985. Histochemistry: Theorethical and Applied. 4th Ed. Churchill, Livingstone, Edinburg, 1055 pp.

Powers M.K., Raymond P.A. 1990. Development of the visual system. In Douglas, R.H., Djamgoz (eds.). The visual system of fish. Chapman and Hall, London, pp. 419-442.

Recassens L., Lombarte A., Morales-Nin B., Torres G. 1998. Spatio-temporal variation in the population structure of the European hake in the NW Mediterranean. J. Fish Biol. 53: 387-401. http://dx.doi.org/10.1111/j.1095-8649.1998.tb00988.x

Recassens L., Chiericoni V., Belcari P. 2008. Spawning pattern and bath fecundity of the European hake (Merluccius merluccius, Linnaeus, 1758) in the western Mediterranean. Sci. Mar. 72(4): 721-732.

Ribeiro L., Sarasquete C., Dinis M.T. 1999. Histological and Histochemical Characteristics during development of the Senegal sole, Solea senegalensis. Aquaculture 171: 291-306.

Roo F.J., Socorro J., Izquierdo M.S., Caballero M.J., Hernández-Cruz C.M., Fernandez A., Fernández-Palacios H. 1999. Development of red porgy, Pagrus pagrus visual system in relation with changes in the digestive tract and larval feeding habits. Aquaculture 179: 499-512. http://dx.doi.org/10.1016/S0044-8486(99)00183-0

Sandy J.M., Blaxter J.M.S. 1980. A study of retinal development in larval herring and sole. J. Mar. Biol. Ass. U.K. 60: 59-71. http://dx.doi.org/10.1017/S0025315400024115

Sánchez P., Sartor P., Recassens L., Ligas A., Martin J., Ranieri S., Demestre M. 2007. Trawl catch composition during different fishing intensity periods in two Mediterranean demersal fishing grounds. Sci. Mar. 71(4): 765-773.

Sánchez F.J., Otero J., Cal J.R., Lago M.J., Gómez C., Iglesias J. 2011. The first spontaneous spawning of European hake, Merluccius merluccius L: Characteristics of eggs and early larval stages. Aqua. Res. (in press).

Sarasquete C., Gutiérrez M. 2005. New Tetrachromic VOF Stain (Type III-G.S) For Normal and Pathological Fish Tissues. European J. Histochem. 49(2): 105-114.

Sarasquete C., Polo A., Pascual E., Yúfera M. 1993a. Histochemistry of proteins, lipids and carbohydrates in the yolk constituents of oocytes, eggs and larvae of Sparus aurata L. Phys. Bioch. Mar. Fish Larv. 309-314.

Sarasquete C., Polo A., González de Canales M.L. 1993b. A histochemical and immunohistochemical study of digestive enzymes and hormones during the larval development of the seabream, Sparus aurata. Histochem. J. 25: 430-437. http://dx.doi.org/10.1007/BF00157807 PMid :7689548

Sarasquete C., Polo A., Yufera M. 1995. Histological and histochemical study during larval development of Sparus aurata L. Aquaculture 130: 79-92. http://dx.doi.org/10.1016/0044-8486(94)00175-N

Sarasquete C., González de Canales M.L., Arellano J., Muñoz-Cueto J.A., Ribeiro L., Dinis M.T. 1996. Histochemical aspects of the yolk-sac and digestive tract of the Senegal sole, Solea senegalensis. Histol. Histopatol. 11: 881-888. PMid :8930630

Sarasquete C., González de Canales M.L, Arellano J.M., Muñoz-Cueto J.A., Ribeiro L., Dinis M.T. 1998. Histochemical study of skin and gills of Senegal sole, Solea senegalensis larvae and adults. Histol. Histopatol. 13: 727-735. PMid :9690130

Sarasquete C., Ribeiro L., Gisbert E., Vieira L., Dinis M.T. 2001. Glycoconjugates in epidermal, branchial and digestive mucous cells and gastric glands of gilthead seabream, Sparus aurata, Senegal sole, Solea senegalensis and Siberian sturgeon, Acipenser baeri. Europ. J. Histochem. 45: 267-278. PMid :11759813

Sarasquete C., Cardenas S., González de Canales M.L., Pascual E. 2002a. Oogenesis in the bluefin tuna, Thunnus thynnus L. A histological and histochemical study. Histol. Histopatol. 17: 775-788. PMid :12168787

Sarasquete C., González de Canales M.L., Piñuela C., Muñoz-Cueto J.A., Rendon C., Mañanos E., Rodríguez-Gómez F.J., Pascual E. 2002b. Histochemical Characteristics of the vitellogenic oocytes of the bluefin, Thunnus thynnus L. Cienc. Mar. 28(4): 419-431.

Segner H., Storch V., Reinecke M., Kloas W., Hamke W. 1994. The development of functional digestive and metabolic organs in turbot, Scophthalmus maximus. Mar. Biol. 119: 35-39. http://dx.doi.org/10.1007/BF00347544

Stroband H.W.J., Dabrowski K.R. 1981. Morphological and physiological aspects of the digestive system and feeding in fresh-water fish larvae. In: Fontaine, M. (ed.), Nutrition des Poissons. Edition de Centre National de la Researche Scientifique, Paris, France, pp: 353-376.

Walzer C., Schönenberger N. 1979. Ultrastructure and cytochemistry study of the yolk syncytial layer in the alevin of trout (Salmo fario trutta L) after hatching. Cell Tissue Res. 196: 59-73. PMid :570459

Zambonino-Infante J., Gisbert E., Sarasquete C., Navarro I., Gutiérrez J., Cahu C.L. 2008. Ontogeny and physiology of the digestive system of marine fish larvae. In: Cyrino, JEO., Bureau, D. and Kapoor, B.G. (eds.), Feeding and Digestive Functions of Fish. Science Publishers, Inc, Enfield (NH), Jersey, Plymouth, pp: 281-348.




How to Cite

Ortiz-Delgado JB, Iglesias J, Sánchez FJ, Cal R, Lago MJ, Otero JJ, Sarasquete C. A morphohistological and histochemical study of hatchery-reared European hake, Merluccius merluccius (Linnaeus, 1758), during the lecitho-exotrophic larval phase. scimar [Internet]. 2012Jun.30 [cited 2022Aug.8];76(2):259-71. Available from: https://scientiamarina.revistas.csic.es/index.php/scientiamarina/article/view/1335




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