Studies on the Physiology of Atractosteus Spatula Larval Development and Its Applications to Early Weaning Onto Artificial Diets
Palabras clave:
alligator gar, larval development, morphology, physiology, enzymes, hormones, nutritionResumen
This paper describes the strategy utilized for the domestication of the alligator gar
(Atractosteus spatula), highlighting different aspects of larval development and the
weaning of larvae onto artificial diets. Developmental stages based on nutritional
characteristics were established. Lecithotrophic stage was observed between 1–4 days after
hatching (DAH) larvae, lecithoexotrophic stage between 5–8 DAH larvae and the
exotrophic stage initiates in 9 DAH larvae. The degree of development for alligator gar
larvae could be differentiated by snout length. In addition, pigmentation could be easily
used to separate individuals in poor condition. Growth rate of alligator gar larvae was 1.55
mm/day during the first 10 DAH and 5.6 mm/day thereafter until 15 DAH. RIA analyses
showed that T3 levels are low during embryonic development, but increase after hatching,
until 10 DAH, decreasing thereafter. The effect of thyroidian hormones and goitrogens on
larval development was evaluated. Snout length was significantly reduced in larvae treated
with thiourea without affecting growth, whereas in larvae treated with T3 snout
development was accelerated. Histological assessment of the development of the digestive
tract revealed that at the onset of exogenous feeding (5DAH) the digestive tract was
completely developed. Measurement of mid-gut cell height was valuable tool for evaluating
the suitability of diets. Additionally, the ontogeny of digestive enzymes was characterized.
Pepsin-like acid proteolytic activity was detected from the beginning of exogenous feeding
(5 DAH). Trypsin, chymiotrypsin and aminopeptidase-like alkaline proteolytic activity
gradually increased between 2-9 DAH. Substrate-gel electrophoresis was utilized to
confirm and further characterize digestive enzyme activity. Growth hormone coding
sequence was obtained by RT-PCR and its expression was detected in embryos and in
several larval stages, revealing an important role during embryogenesis and organogenesis.
RNA/DNA ratios were determined and proven to be a useful indicator of larval nutritional
condition. In fed larvae RNA/DNA ratio rose until the 13 DAH, while in starved larvae the
ratio RNA/DNA was low and remained constant. Three feeding experiments were carried
out and some important characteristics of artificial diets were identified, namely floatability
and size. The results of the feeding trials made it possible to culture larvae and juveniles
using exclusively artificial diets.
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Adelman, I. A., 1977. Effect of bovine growth hormone on the growth of carp (Cyprinus carpio) and the
influences of temperature and photoperiod. J. Fish. Res. Bd. Can. 34,509-515.1977
Agellon, L. B., Emery, C. J., Jones, J. M., Davies, S. L., Dingle, A. D., Chen, T. T., 1988. Promotion of rapid
growth of rainbow trout (Salmo gairdnieri) by a recombinant fish growth hormone. Can. J. Fish.
Aquat. Sci., 45, 146-151.
Aguilera, C., 1999. Bases fisiológicas del desarrollo de larvas de catán (Atractosteus spatula) y perspectivas
para su cultivo. Tesis Doctoral, Facultad de Ciencias Biológicas, U. A. N. L. Monterrey, N. L.,
pp.186.
Aguilera, C., Mendoza, R., Rodríguez, G., Márquez, G., 2002. Morphological description of alligator gar and
tropical gar larvae, with an emphasis on growth indicators. Trans. Amer. Fish. Soc., 131, 899-909.
Alliot, E., Pastoureaud, A., Trellu, J., 1977. Evolution des activités enzymatiques dans le tube digestif au
cours de la vie larvaire du bar (Dicentrarchus labrax). Variations des proteinogrammes et des
zymogrammes. Actes de Colloques du C.N.E.X.O., 4, 85-91.
Alvarez, J., Sarmiento, M., Mendoza, R., 1999. Estudio histológico del desarrollo del tracto digestivo en
larvas de catán Atractosteus spatula en condiciones de inanición. Resúmenes del XXIII Congreso
Nacional de Histología, Monterrey, México, pp. 35.
Appelbaum, S., 1985. Rearing of the dover sole, Solea solea (L), through its larval stages using artificial diets.
Aquaculture 49, 209-221.
AQUACOP, Calvas, J., 1990. State of the art of IFREMER in tropical aquaculture. In: Advances in tropical
Aquaculture, Aquacop-IFREMER, Actes de Colloque 9, pp 625-642
Baragi, V., Lovell, R. T., 1986. Digestive enzyme activities in striped bass from first feeding through larval
development. Trans. Am. Fish. Soc. 115, 478-484.
Beccaria, C., Diaz, J. P., Connes, R., Chatain, B., 1991. Organogenesis of the exocrine pancreas in the sea
bass, Dicentrarchus labrax L., reared extensively and intensively. Aquaculture 99, 339-354.
Bromley, P. J., 1978. The weaning of hatchery reared turbot larvae (Scophtalmus maximus L.) on a dry diet.
Aquaculture 13, 339-345.
Brown, C. L., Sullivan, C. V., Bern, H. A., Dickhoff, W. W., 1987, Occurrence of thyroid hormones in early
developmental stages of teleost fish. American Fisheries Society Symposium, 2, 144-150.
Buckley, L., Caldarone, E., Ong, T.-L., 1999. RNA-DNA ratio and other nucleic acid-based indicators for
growth and condition of marine fishes. Hydrobiologia. 401, 265-277.
Buddington, R. K., 1985. Digestive secretions of lake sturgeon Acipenser fulvescens during early
development. J. Fish. Biol., 26, 715-724. 341
Buddington, R. K., Chrstofferson, P., 1985. Digestive and feeding characteristics of the chondrosteans. In:
North american sturgeons: biology and aquaculture potential, (ed. by F.P. Binkowski and S.I.
Doroshov), DR W. Junk Publishers, 31-41. Kluwer Academic Publishers Group, Boston.
Buddington, R. K., Dorshov, S. I., 1986. Development of digestive secretions in white sturgeon juveniles
(Acipenser transmontanus) Comp. Biochem. Physiol., 83A,233-238. 344
Burns, T. A., Stalling, D. T., Goodger, W., 1981. Gar ichthyotoxin, its effect on crayfish, with notes on
bluegill sunfish. The Southwestern Naturalist 25(4), 513-515.
Busch, A., 1996. Transition from endogenous to exogenous nutrition: larval size parameters determining the
start of external feeding and size of prey ingested by ruegen spring herring Clupea harengus. Marine
Ecology Progress Series 130, 39-46.
Chavez-Lomelí, M. O., 1989. Biología de los peces del río San Pedro en vista de determinar su potencial para
piscicultura. Instituto de Investigaciones sobre Recursos Bióticos y la Fundación Universitaria para
la cooperación, Jalapa, Ver. 90 pp.
Cheema, I. R., Maty, A. J., 1978. Increased uptake of L-leucine-14C in the skeletal muscle of rainbow trout,
Salmo gairdnieri after administration of growth hormone. Pak. J. Zool. 10,119-123.
Clemmesen, C., 1996. Importance and limits of RNA/DNA ratios as a measure of nutritional condition in fish
larvae. In: Survival strategies in early life stages of marine resources (ed. by Watanabe, Y.,
Yamashita, Y. and Oozeki, Y.) Proceedings of an Internatíonal Workshop, Yokohama, Japan, 11-14
October 1994. AA. Balkema, Rotterdam, the Netherlands, pp. 67-82
Cook H., Cook, A. F., Peter, R. E., 1983. Ultrastructural immuno-chemistry of growth hormone cells in the
goldfish pituitary gland. Gen. Comp. Endocrinol., 50,348-353.
Dabrowski, K. R., 1986. Ontogenical aspects of nutritional requerimients in fish. Comp. Biochem. Physiol.,
A(4),639-655.
Dabrowski, K., Bardega, R., 1984. Mouth size and predicted food size preferences of larvae of three cyprinid
fish species. Aquaculture 40, 41-46.
Dabrowski, K., Culver, D., 1991. The physiology of larval fish, digestive tract and formulation of starter diets.
Aquaculture Magazine March/April: 49-61.
Dabrowski, K., Dabrowska, H., Grudniewski, C., 1978. A study of the feeding of common carp larvae with
artificial food. Aquaculture 13, 257-264.
Dabrowski, K., Kaushik, S. J., Fauconneau, B., 1985. Rearing of sturgeon (Acipenser baeri Brandt) larvae I.
Feeding Trial. Aquaculture 47, 185-192.
Dean, B., 1895. The early development of gar-pike and sturgeon. Journal of Morphology 11(1), 1-55.
Dettlaff, T. A., Ginsburg, A. S., Schmalhausen, O. I., 1993. Development of prelarvae. In: Sturgeon Fishes,
Developmental Biology and Aquaculture. Pp. 155-221 Springer-Verlag Ed., Berlin, Germany.
Devresse, B., Candreva, P., Sorgeloos, P., 1991. A new artificial diet for the early weaning of seabass
(Dicentrarchus labrax) larvae. In: Larvi 91 - Fish and crustacean larviculture symposium, European
Aquaculture Society, Special Publication, 15, 178-182.
Duray, M., Bagarinao, T., 1984. Weaning of hatchery-bred milkfish larvae from live food to artificial diets.
Aquaculture 325-332.
Echelle, A. A., Riggs, C. D., 1972. Aspects of the early life history of gars (Lepisosteus) in Lake Taxoma.
Transactions of the American Fisheries Society 101, 106-112.
FAO/PNUMA, 1984. Conservación de los recursos genéticos de los peces: Problemas y recomendaciones.
Informe de la consulta de expertos sobre los recursos genéticos de los peces. FAO. Doc. Tec. Pesca,
(217): 42p.
Ferraris, R. P., Tan, J. D., De La Cruz, M. C., 1987. Development of digestive tract of milkfish, Chanos
chanos (Forsskal): hitology and histochemistry. Aquaculture 61, 241-257.
Ferron, A., Leggett, W. C., 1994. An appraisal of condition measures for marine fish larvae. Adv. Mar. Biol.
, 217-303.
Fauconneau, B., Mady, M. P., LeBail, P. Y., 1996. Effect of growth hormone on muscle protein synthesis in
rainbow trout (Oncorhynchus mykiss) and Atlantic salmon (Salmo salar).Fish Physiol. Biochem.
(1), 49-56.
Foster, A. R., Houlihan, D. F., Gray, C., Medale, F., Fauconneau, B., Kaushik, S. J., LeBail, P. Y., 1991. The
effects of ovine growth hormone on protein turnover in rainbow trout. Gen. Comp. Endocrinol. 81,
-120.
Fuchs, J., Nedelec, G., 1990. Larvae rearing and weaning of seabass, Lates calcarifer (Block), on
experimental compounded diets. Advances in Tropical Aquaculture, Actes de Colloque 9 pp. 677-
Fukuda, M., Sako, H., Shigeta, T., Shibata, R., 2001. Relationship between grow and biochemical indices in
laboratory-reared juvenile Japanese flounder (Paralichthys olivaceus), and its application to wild
fish, Mar. Biol., 138, 47-55.
García, J., Márquez, G., Páramo, S., 1997. Utilización de alimento fresco y artificial en la sustitución de
alimento vivo para la cría de larvas y postlarvas del pejelagarto Atractosteus tropicus. In: Memorias
de la Semana de Divulgación y Video Científico 1997, Secretaría de servicios académicos
Universidad Juárez Autónoma de Tabasco, pp. 61-64.
Gatsoupe, F. J., Luquet, P., 1982. Weaning of the sole (Solea solea) before metamorphosis. Aquaculture,
:359-368.
Gawlicka, A., Teh, S. J., Hung, S. S. O., Hinton, D. E., de la Noue, J., 1995. Histological and histochemical
changes in the digestive tract of white sturgeon larvae during ontogeny. Fish Physiol. Biochem., 14,
-371
Gershanovich, A. D., 1991. Lipid mobilization during early development of sturgeons. Ed. Acipenser,
Cemagref Publ. pp. 41-51. P. Williot, Ed. Acipenser, Cemagref Pub., Moscow.
Gill, J. A., Sumpter, J. P., Donaldson, E. M., Dye, H. M., Souza, L., Berg, T., Wypych, J., Langley, K., 1985.
Recombinant chicken and bovine growth hormones accelerate growth in aquacultured juvenile
Pacific salmon, Oncorhynchus kisutch. Biotechnology 3, 643-646.
Gisbert, E., Rodríguez, A., Castelló-Orvay, F., Williot, P., 1998. A histological study of the development of
the digestive tract of siberian sturgeon (Acipenser baeri) during early ontogeny. Aquaculture 167,
-209.
Goodyear, C. P., 1967. Feeding Habits of Three Species of Gars, Lepisosteus, along the Mississippi Gulf
Coast. Trans. Am. Fish. Soc. 96, 297-300
Govoni, J. J., George, W., Watanabe, Y., 1986. The physiology of digestion in fish larvae. Environmental
Biology of Fishes, 16 (1-3), 59-77.
Grant, G. C., 1996. RNA-DNA ratios in white muscle tissue biopsies reflect recent growth rates of adult
brown trout. J. Fish Biol. 48, 1223-1230.
Grizzle, J. M., Curd, M. R., 1978. Posthatching histological development of the digestive system and swim
bladder of logperch, Percina caprodes. Copeia, 3, 448-455.
Hecht, T., Pienaar, A. G., 1993. A review of cannibalism and its implications in fish larviculture. Journal of
the World Aquaculture Society 24(2), 246-261.
Hernández, U., 2002. Control reproductivo de Atractosteus tropicus: Evaluación de la inducción hormonal.
Tesis de Maestría. Facultad de Ciencias Biológicas, U. A. N. L., Monterrey, N. L., 98 p
Hernandez, U., Marquez, G., Páramo, S., Félix, S., Hernández, S., 1997. Valor nutritivo de nauplios de
Artemia spp. y uso en la larvicultura del pejelagarto Atractosteus tropicus. In: Memorias de la
Semana de Investigación y Divulgación Científica de la Universidad Juárez Autónoma de Tabasco
pp. 65-68.
Hernández, U., 1999. Punto crítico de no-retorno en larvas de pejelagarto Atractosteus spatula (Gill, 1863).
Tesis de Licenciatura Inédita. División Académica de Ciencias Biológicas, Universidad Juárez
Autónoma de Tabasco 1997. pp. 46
Higgs, D. A., Donaldson, E. M., Dye, H. M., McBride, JR., 1975. A preliminary investigation of the effect of
bovine growth hormone on growth and muscle composition of coho salmon (Oncorhynchus kisutch).
Gen Comp. Endocrinol., 27, 240-253.
Higgs, D. A., Donaldson, E. M., Dye, H. M., McBride, JR., 1976. Influence of bovine growth hormone and Lthyroxine
on growth, muscle composition and histological structure of the gonads, thyroid, pancreas
and pituitary of coho salmon (Oncorhynchus kisutch). J. Fish. Res. Board Ca., 33, 1585-1603.
Holloway, A., 1954. Notes on the life history and management of the shortnose and longnose gars in Florida
waters. Journal of Wildlife Management 18(4), 438-449
Houlihan, D. F., Mathers, E. M., Foster, A., 1993. Biochemical correlates of growth rate in fish. In: Fish
Ecophysiology (ed. by J. Cliff and B. Jensen) pp. 45-71. Chapman and Hall, London. pp.421
Hunter, J. R., 1981. Feeding Ecology and Predation of Marine Fish Larvae. In: Marine Fish Larvae (ed. by
Lasker, R.), pp. 34-59. Washington Sea Grant Program Publisher.
Inui, Y., Tagawa, M., Miwa, S., Hirano, T., 1989. Effects of bovine TSH on the tissue thyroxine level and
metamorphosis in prometamorphic flounder larvae. General and Comparative Endocrinology, 74,
-410.
Kaji, T., Tanaka, M., Takahashi, Y., Oka, M., Ishibashi, N., 1996. Preliminary observations on development
of pacific bluefin tuna Thunnus thynnus (Scombridae) larvae reared in the laboratory, with special
reference to the digestive system. Mar. Freshwater Res., 4, 261-269.
Kobuke, L., Specker, J., Berni, H. A., 1987. Thyroxine content of eggs and larvae of coho salmon
Oncorhynchus kisutch. J. Exp. Zool. 248, 168-176
Kolkovski, S., Tandler, A., Kissil, G. W., Gertler, A., 1993. The effect of dietary exogenus digestive enzymes
on ingestion, assimilation, growth and survival of gilthead seabream (Sparus aurata, Sparidae,
Linnaeus) larvae. Fish Physiology and Biochemistry, 12(3), 203-209.
Komourdjian, M. P., Saunders, R. L, Fenwick, J. C., 1976. The effect of porcine somatotropin on growth and
survival in seawater of Atlantic salmon (Salmo salar). Can. J. Zool. 54, 531-535.
Lacroix, M. C., Devinoy, E., Cassy, S., Servely, J. L., Vidaud, M., Kann, G., 1999. Expression of growth
hormone and its receptor in the placental and feto-maternal environment during early pregnancy in
sheep. Endocrinology, 140(12), 5587-97.
Lam, T. J., 1980. Tyroxine enhances larval development and survival in Sarotherodon (Tilapia) mossambicus
Rupell. Aquaculture: 21, 287-291.
Lauf, M., Hoffer, R., 1984. Proteolytic enzymes in fish development and the importance of dietary enzymes.
Aquaculture 37, 335-346
Lazo, J., 2000. Conocimiento actual y nuevas perspectivas en el desarrollo de dietas para larvas de peces
marinos. In: Avances en nutrición acuícola V. Memorias del 5º Simposium Internacional de
Nutrición acuícola (ed. by Cruz- Suárez, L. E., Ricque-Marie, O., Tapia-Salazar, M., Olvera-Novoa,
M. A. y Civera-Cerecedo, R.), pp. 300-312. 19-22 Noviembre, 200. Mérida, Yuc., México
León, L., Aguiar, R., Hernández, 1978. Estudio sobre la biología y el cultivo artificial del manjuari
(Atractosteus tristoechus) Bloch y Schmeider. Ministerio de la Industria Pesquera, Dirección Ramal
de Acuicultura, Investigación No. 85, Cuba. Pp. 1-35.
Lovshin, L. L., Rushing, H. J., 1989. Acceptance by largemouth bass fingerlings of pelleted feeds with a
gustatory additive. The Progressive Fish-Culturist, 51, 73-78.
Maldonado, E., 1991. Aprovechamiento de peces forrajeros en la alimentación de Atractosteus tropicus (Gill)
en jaulas flotantes en el Estado de Tabasco, Mexico. Universidad y Ciencia, 8(15), 77-90.
Markert, J. R., Higgs, D. A., Dye, H. M., MacQuarrie, D. W., 1977. Influence of bovine growth hormone on
growth rate, appetite, and food conversion of yearling coho salmon (Oncorhynchus kisutch) fed two
diets of different composition. Can. J. Zool., 55, 74-83.
Márquez, G., 2000. Biología y Tecnología para el Cultivo del pejelagarto en el sureste de México. In: Redes
nacionales de investigación en acuacultura, Memorias de la IV Reunión. (ed. by P. Álvarez, M.
Guzmán, S. Contreras, y A. Silva), pp. 263-265. SEMARNAP, Instituto Nacional de la Pesca.
México.
Marr, C., 1999. Improving efficiency in starter feeds. Aqua Feed, 4, 18-20.
Mathers, E. M., Houlihan, D. F., Burren, L. J., 1994. RNA, DNA and protein concentrations in fed and
starved herring Clupea harengus larvae. Mar Ecol. Prog. Ser. 107, 223-231.
Matty, A. J., 1962. Effects of mammalian growth hormone on Cottus scorpius blood. Nature (London). 195,
-507.
May, E. B., Echelle, A. A., 1968. Young-of-year alligator gar in Lake Taxoma, Oklahoma. Copeia. 3, 629-
McFadzen, I. R., Lowe, D. M., Coombs, S. H., 1994. Histological changes in starved turbot larvae
(Schophtalmus maximus) quantified by digital image analysis. Journal of Fish Biology, 44, 255-262.
Medale, F., Fauconneau, B., Kaushick, S. J., 1988. Effect of GH administration on energy metabolism in
rainbow trout. In: Energy Transformation in Cells and Organisms (ed. by W. Wieser and E.
Gnaiger), pp. 240. Georg Thieme Verlag, Stuttgart.
Mendoza, R., 2000. Catán. In: Estado de salud de la acuacultura, (ed. by Alvarez-Torres), pp. 231-242.
Instituto Nacional de la Pesca, Secretaria de Recursos Naturales y Pesca. Distrito Federal, Mexico.
Mendoza, R., Aguilera, C., 2000a. Morphophysiological studies on alligator gar Atractosteus spatula larval
development as a basis of their weaning onto artificial diets. 24th Annual Larval Fish Conference.
Alabama.
Mendoza, R., Aguilera, C., 2000b. Cultivo y pesquería de catán, In: Carta Nacional Pesquera pp. 56-57..
Diario Oficial de la Federación, Tomo DLXIII, No. 20, 28 de agosto de 2000.
Mendoza, R., Aguilera, C., Montemayor, J., Rodríguez, G., Márquez, G., 2000a. Biología de los lepisosteidos
y estudios orientados hacia la recuperación de las poblaciones naturales del catán (Atractosteus
spatula) (Lacepède, 1803). In: Redes nacionales de investigación en acuacultura, Memorias de la IV
Reunión P. (ed. by Álvarez, Guzmán, M., Contreras, S. y Silva, A.), pp. 103-120. SEMARNAP,
Instituto Nacional de la Pesca. Distrito Federal, México.
Mendoza, R., Aguilera, C., Montemayor, J., Rodríguez, G., Castro, R., 2000b. Aspectos reproductivos de los
lepisosteidos, con énfasis en las perspectivas para el control de la reproducción del catán
(Atractosteus spatula). In: Redes nacionales de investigación en acuacultura, Memorias de la IV
Reunión P. (ed. by Álvarez, Guzmán, M., Contreras, S. y Silva, A.), pp 281-290. Redes nacionales
de investigación en acuacultura, Memorias de la IV Reunión. SEMARNAP, Instituto Nacional de la
Pesca. Distrito Federal, México.
Mendoza, R., Aguilera, C., Rodríguez, G., Márquez, G., 2000c. Estrategias para la domesticación de especies
en acuacultura: El catán (Atractosteus spatula). In: Redes Nacionales de Investigación en
Acuacultura, Memorias de la V Reunion (ed. by Alvarez Torres, Ramírez-Flores, Torres-Rodriguez
y Mora-Cervantes), pp. 95-102. Instituto Nacional de Pesca-SEMARNAP. Distrito Federal, México.
Mendoza, R., Aguilera, C., Ruiz, V., García-Carreño, F., Nolasco, H., Montemayor J., Rodríguez, G., 2000d.
Enzyme ontogenetic larval development of the alligator gar Atractosteus spatula in México:
Perspectives for its culture. World Aquaculture Society. New Orleans, USA. Feb, 2000. page 219.
Mendoza, R., Aguilera, C., 2001. Bases Fisiológicas del desarrollo de larvas de Atractosteus spatula y
perspectivas para su cultivo. Ciencia-UANL. 4(2), 161-166
Mendoza, R., Aguilera, C., Montemayor, J., 2001. Effects of thyroid hormones and glucocorticoids on
alligator gar Atractosteus spatula larval development. World Aquaculture Society. Orlando, Florida,
USA. Page 434.
Mendoza, R., Aguilera, C., Rodriguez, G., González, M., Castro, R., 2002a. Morphophysiological studies on
alligator gar (Atractosteus spatula) larval development as a basis for their culture and repopulation of
their natural habitats. Reviews in Fish Biology and Fisheries. 00: 1-9, In press.
Mendoza, R., Aguilera, C., Carreón, L., Montemayor, J., González, M., 2002b. Early weaning of alligator gar
Atractosteus spatula larvae. Aquaculture 2002, WAS. San Diego, California, Enero 27-30 2002.
Page 214.
Mendoza, R., Aguilera, C., Carreón, L., 2002c. Acidos nucleicos para evaluar la condición de larvas de peces.
Ciencia-UANL, 5(2), 211-217.
Mendoza, R., Garza-Rodriguez, M. L, Hernández-Montenegro, V., Aguilera, C., Barrera Saldaña, H., Revol,
A., 2002d. Cloning of the growth hormone cDNA of alligator gar Atractosteus spatula and its
expression along larvae stages. World Aquaculture Society. San Diego, California USA. January,
Page 213.
Mommsen, T., 1998. Growth and Metabolism. In: The Physiology of Fishes. (ed. by D.H., Evans). CRC
Press, Boca Raton, Florida. pp. 65-97.
Moore, G., Trautman, M., Curd, M., 1973. Description of postlarval gar (Lepisosteus spatula Lacepede,
Lepisosteidae), with a list of associated species from the Red River, Choctaw County, Oklahoma.
The Southwest Naturalist 18, 343-344.
Moyano, F. J., Diaz, M., Alarcon, F. J., Sarasquete, M. C., 1996. Characterization of digestive enzyme
activity during larval development of gilthead sea bream (Sparus aurata). Fish Physiol. Biochem.,
, 121-130.
Munilla-Moran, R., Saborido, F., 1996. Digestive enzymes in marine species I. Proteinases activities in gut
from red fish (Sebastes mentella), seabream (Sparus aurata) and turbot (Scophtalmus maximus).
Comp. Biochem. Physiol. 113B, 395- 402. 384
Netch, N. F., Witt, A., 1962. Contributions to the life history of the longnose gar (Lepisosteus osseus) in
Missouri. Transactions of the American Fisheries Society 91(3), 251-262.
Oozeki, Y., Ishii, T., Hirano, R., 1989. Histological study of the effects of starvation on reared and wildcaught
larval stone flounder Kareius bicoloratus. Mar. Biol. 100, 269-275
Pantaleon, M., Whiteside, E. J, Harvey, M. B, Barnard, R. T, Waters, M. J., Kaye, P. L., 1997. Functional
growth hormone (GH) receptors and GH are expressed by preimplantation mouse embryos: a role for
GH in early embryogenesis?. Proc Natl Acad Sci U S A., 94(10), 5125-30.
Pearson, W. D., Thomas, G. A., Clark, A. L., 1979. Early piscivory and timing of the critical period in
postlarval longnose gar at mile 571 of the Ohio River. Transactions of the Kentucky Academy of
Science 40(3-4), 122-128.
Perez-Sanchez, E., 1995. Efecto de la gonadotropina coriónica humana (GCH) en la maduración gonádica del
pejelagarto (Atractosteus tropicus) Gill 1823, en condiciones de laboratorio. Tesis de licenciatura
Inédita, Universidad Juárez Autónoma de Tabasco, División Académica de Ciencias Biológicas, 28
pp.
Person-Le Rouyet, J., 1990. Early weaning of fish larvae onto microdiets: constraints and perspectives. In:
Advances in tropical Aquaculture, Aquacop-IFREMER, Actes de Colloque 9, pp 625-642.
Pickford, G. E., Atz, J. W., 1957. The Physiology of the Pituitary Gland of Fishes, pp. 1-91. New York
Zoological Society, New York.
Pickford, G. E., Thompson, E. F., 1948. The effects of purified mammalian growth hormone on the killfish
Fundulus heteroclitus (Linn). J. Exp. Zool. 109, 367-383
Resendez, A., Salvadores, M. L., 1983. Contribución al conocimiento de la biología del pejelagarto
Lepisosteus tropicus (Gill) y la tenguayaca Petenia splendida (Günther), del Estado de Tabasco.
Biotica 8(4), 413-426.
Rodríguez, F., Marquez, G., Paramo, S., 1997. Evaluación del uso de cladóceros y nauplios de Artemia en la
alimentación de larvas de pejelagarto Atractosteus tropicus. In: Memorias de la Semana de
Investigación y Divulgación Científica . pp. 72-75. Universidad Juárez Autónoma de Tabasco 1997.
Rodríguez, G., Mendoza, R., Aguilera, C., Montemayor, J., 2000. Elaboración de una dieta artificial para
larvas de catán (Atractosteus spatula) con miras a su repoblación y posible cultivo. In: Investigación
para el Desarrollo Regional. pp. 160 -164Consejo Nacional de Ciencia y Tecnología. Ed., Distrito
Federal, Mexico.
Rojas, P., Mendoza, R., 2000. El cultivo de especies natives en México. In: Estado de salud de la acuacultura
(ed. by .. Alvarez-Torres, P.). pp. 431-476. Instituto Nacional de la Pesca, Secretaria de Recursos
Naturales y Pesca. Distrito Federal, Mexico.
Rooker, J., 1997. Early life history of Red drum (Sciaenops ocellatus) in subtropical seagrass meadows:
patterns of condition, growth, and mortality. Ph.D. Dissertation. Faculty of the Graduate School of
the University of Texas at Austin. Pp. 191.
Rosas, M., 1976. Peces dulceacuícolas que se explotan en México y datos sobre su biología. Instituto
Nacional de la Pesca/Subsecretaria de Pesca. México. Pp. 90.
Rosca, R., Appelbaum, S., 1985. Experiments on the suitability of dry food for larvae of Coregonus lavaretus
L. Aquaculture, 48, 291- 302
Rubin, D. A., Youson, J. H., Marra, L. E., Dores, R. M., 1996. Cloning of a gar (Lepisosteus osseus) GH
cDNA: trends in actinopterygian GH structure. J. Mol. Endocrinol. 16 (1), 73-80
Ruiz, V., 2000. Caracterización bioquímica de las principales enzimas digestivas del catán (Atractosteus
spatula). Tesis de Licenciatura, Facultad de Ciencias Biológicas, U. A. N. L. Monterrey, N. L., 76 pp
Sarasquete, M. C., Polo, A., Yufera, M., 1995. Histology and histochemistry of the development of the
digestive system of larval gilthead seabream, Sparus aurata L. Aquaculture, 130, 79-92.
Segner, H., Stroch, V., Reinecke, M., Kloas, W., Hanke, W., 1994. The development of functional digestive
and metabolic organs in turbot, Scophtalmus maximus. Marine Biology, 119, 471-486.
SEPESCA-INP (Secretaria de Pesca - Instituto Nacional de la Pesca), 1994. Atlas Pesquero de México.
SEPESCA. México. Ed. by M. Rodriguez, M. Palacios, R. Cruz and C. Díaz Pp. 234.
Sheridan, M. A., 1986. Effects of thryroxine, cortisol, growth hormone and prolactin on lipid metabolism of
coho salmon Oncorhynchus kisutch, during smoltification. Gen. Comp. Endocrinol. 64, 220-238.
Simon, T. P., Tyberghein, E. J., 1991. Contributions to the early life history of the spotted gar, Lepisosteus
oculatus Winchell, from Hatchet Creek, Alabama. Transactions of the Kentucky Academy of
Science 52(3-4), 124-131.
Simon, T. P., Wallus, R., 1989. Contributions to the early life histories of gar (Actinopterygii: Lepisosteidae)
in the Ohio and Tennessee River basins with emphasis on larval development. Transactions of the
Kentucky Academy of Science 50(1-2), 59-74.
Sullivan, C. V., Iwamoto, R. N., Dickhoff, W. W., 1987. Thyroid hormones in blood plasma of developing
salmon embryos. General and Comparative Endocrinology. 65, 337-345.
Suttkus, R. D., 1963. Order Lepisostei. In: Fishes of the Western North Atlantic. Part three, Soft-rayed fishes
(ed. by . H. B. Bigelow) pp. 61-88. Yale University Memoir Sears Foundation for Marine Research,
New Haven, Connecticut.
Tagawa, M., Hirano, T., 1987. Effects of thyroid deficiency in eggs on early development of the medaka,
Oryzias latipes. The Journal of Experimental Zoology, 257, 36-366.
Tagawa, M., Tanaka, M., Matsumoto, S., Hirano, T., 1990. Thyroid hormones in eggs of various freshwater,
marine and diadromus teleosts and their changes during egg development. Fish Physiology and
Biochemistry. 8(6), 515-520.
Tanaka, M., 1969a. Studies on the structure and function of the digestive system in teleost larvae I.
Development of the digestive system during prelarval stage. Japanese Journal of Ichthyology, 16(1),
-8.
Tanaka, M., 1969b. Studies on the structure and function of the digestive system in teleost larvae II.
Characteristic of the digestive system in larvae at the stage of first feeding. Japanese Journal of
Ichthyology, 16(2), 41-49.
Tanaka, M., 1971. Studies on the structure and function of the digestive system in teleost larvae III.
Development of the digestive system during postlarval stage. Japanese Journal of Ichthyology, 18(4),
-173.
Tanaka, M., 1972. Studies on the structure and function of the digestive system in teleost larvae V. Epithelial
changes in the posterior gut and protein ingestion. Japanese Journal of Ichthyology, 19(3), 172-180.
Tandler, A., Kolkovski, S., 1991. Rates of ingestion and digestibility as limiting factors in the successful use
of microdiets in Sparus aurata larval rearing. European Aquaculture Society, Special Publication,
, 169-171.
Theilacker, G. H., Porter, S. M., 1995. Condition of larval walleye pollock, Theragra chalcogramma, in the
western Gulf of Alaska assessed with histological and shrinkage indices. Fish. Bull., 93, 333-344.
Theilacker, G. H., Watanabe, Y., 1989. Midgut cell height defines nutritional status of laboratory raised larval
northern anchovy, Engraulis mordax. Fish. Bull. 87(3), 457-469
Ueberschär, B. F. R., 1988. Determination of the nutritional condition of individual marine fish larvae by
analyzing g their proteolytic enzyme activities with a highly sensitive fluorescence technique.
Meeresforsch. Rep. mar. Res. 32, 144-154.
Verreth, J., Van Tongeren, T., 1989. Weaning time in Clarias gariepinus (Burchell) larvae. Aquaculture, 83,
-88
Verreth, J. A., Torreele, E., Spazier, E., Sluiszen, A. V., 1992. The development of a functional digestive
system in the african catfish Clarias garipinus (Burchell). J. World Aquaculture Soc. 23(4), 286-298
Vonk, H. J., Western, J., 1984. Factors influencing enzymatic digestion. In: Comparative Biochemistry and
Physiology of Enzymatic digestion. Academic Press, pp 448-491
Wagner, C. F., McKeown, B. A., 1981. The purification, partial characterization and bioassay of growth
hormone from two species of Pacific salmon. In: B. Lofts and W.N. Holmes (editors), Current trends
in Comparative Endocrinology. Proc. 9th Symp. Comp. Endocrinol. Hong Kong,p. 211.
Walford, J., Lam, T. J., 1993. Development of digestive tract and proteolytic enzyme activity in seabass
(Lates calcarifer) larvae and juveniles. Aquaculture, 109, 187-205.
Watanabe, T., Kiron, V., 1994. Prospects in larval fish dietetics. Aquaculture, 124, 223-251.
Watanabe, Y., 1984. Morphological and functional changes in rectal epithelium cells of pond smelt during
postembrionic development. Bulletin of the Japanese Society of Scientific Fisheries, 50(5), 805-814.
Westerman, M. E., Holt, G. J., 1988. The RNA-DNA ratios: measurement of nucleic acids in larval Sciaenops
oceIlatus. Contributions to Marine Sciences 30, 117-124.
Xu, K., Wei, Y., Guo, L., Zhu, Z., 1991. The effects of growth enhancement of human growth hormone gene
transfer and human growth hormone administration on crussian carp (Carasius auratus gibelio
Bloch). Acta Hydrobiol. Sin. 15, 31-34.
Yang, B., Green, M., Chen, T., 1999. Early Embryonic Expression of the growth hormone family Protein
Genes in the Developing Rainbow Trout, Onchorhynchus mykiss. Mol Reprod. Dev. 53(2), 127-34.
Yeager, B. L., Bryant, R. T., l983. Larvae of the longnose gar, Lepisosteus osseus, from the Little River in
Tennessee. Journal of the Tennessee Academy of Science 58(1-2), 20-22.
Zitzow, R. E., Millard, J. L., 1988. Survival and growth of lake whitefish (Coregonus clupeaformis) larvae
fed only formulated diets. Aquaculture, 69, 105-113.
