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Sexado 415

SELECCIN DEL SEXO EN MAMFEROS

Ligia M Cantarelli Pegoraro


Maria Gabriela T Rheingantz
& Gustavo A Palma

Introduccin

El origen de la determinacin sexual de la descendencia de los mamferos siempre fue motivo


de especulaciones. Hace ms de 3000 aos, varias teoras, suposiciones y creencias fueron
postuladas. En la Grecia antigua los filsofos meditaban sobre la teora de la determinacin sexual,
segn el lado izquierdo o derecho y del estado caliente o fro. Se especulaba que el testculo del lado
derecho originaba descendencia macho y el lado izquierdo hembras. El lado del tero materno fue
considerado tambin un factor determinante. La posicin durante el acto sexual (norte/sur) favoreca
la generacin de machos y hembras. Los estudios cientficos que colaboraron con la dilucidacin de
los factores involucrados en la determinacin sexual comenzaron efectivamente con el
descubrimiento del espermatozoide, en el siglo XVII, su origen y funcin en el ao 1841, con el
descubrimiento de la gameta femenina en el ao 1827 y con la constatacin de la ocurrencia de la
fecundacin, solamente al final del mismo siglo. El misterio de la generacin fue dilucidado al final del
siglo XIX con el descubrimiento de los cromosomas. En esta poca se consideraba tambin que las
condiciones ambientales eran determinantes para el sexo de los productos, que las condiciones
nutricionales adversas originaban machos, mientras que las favorables hembras. As como fue
relatado que las futuras madres que se alimentaban con dietas basadas en alimentos amargos
generaban hijos de sexo masculino. A inicios del siglo XX, con el descubrimiento de los cromosomas
humanos y de otros mamferos se estableci la importancia de los cromosomas sexuales X e Y,
siendo el Y responsable por la generacin de productos del sexo masculino (ver revisin sobre
mitologa de la determinacin del sexo de Beatty, 1974, 1978; Gledhill, 1988a, b; Mittwoch, 2005).
En mamferos euterianos se determin que la diferenciacin sexual es un proceso secuencial
que se inicia con el establecimiento del sexo gentico o cromosmico (cromosomal) en el momento
de la fecundacin (XX o XY). El espermatozoide portador del cromosoma Y determinar la formacin
de un individuo del sexo masculino y el espermatozoide portador de X uno de sexo femenino. La
primera fase de la diferenciacin sexual es denominada determinacin primaria del sexo y es cuando
tiene lugar la organognesis gonadal, en funcin del sexo gentico del embrin, culminando con la
formacin del fenotipo sexual. Cada etapa de este proceso es dependiente de la etapa precedente y,
en circunstancias normales, el sexo cromosmico est de acuerdo con el sexo fenotpico (Jost y col.
1973; George y Wilson, 1994).

Sexo Sexo Sexo


Cromosmico Gonadal Fenotpico

Fig 1. Esquema de la determinacin sexual en los mamferos

Tanto el macho (XY) como la hembra (XX) poseen gnadas idnticas en la fase inicial del
desarrollo, cuando la cresta gonadal es denominada gnada indiferenciada o gnada totipotente. La
segunda fase de la diferenciacin sexual abarca el desarrollo de la gnada indiferenciada en
testculos y ovarios. La tercera fase es la diferenciacin de los conductos que consiste en la formacin
de los conductos de Wolf en el embrin macho y de los conductos de Mller en embrin de sexo
femenino. Dos hormonas secretadas por el testculo, antimlleriana (AMH) y testosterona,
transforman entonces el tracto urogenital indiferenciado en tracto genital caracterstico del macho. La
hormona antimlleriana es secretada por las clulas de Sertoli y causa regresin de los conductos
416 Pegoraro, Rheingantz & Palma
femeninos (de Mller) en el embrin masculino. La testosterona, secretada por las clulas de Leydig
fetales es responsable del restante desarrollo del tracto reproductivo masculino, incluido la
estabilizacin y diferenciacin de los conductos de Wolf en los rganos secundarios masculinos y la
formacin de los genitales externos y prstata. La ltima fase es la diferenciacin del sinus urogenital
y de los genitales externos, de acuerdo con el sexo del producto (Josso y col. 1977; Baskur, 1980).
En los mamferos euterianos, el cromosoma X es grande, representa 5% del genoma haploide y
es portador de genes que codifican numerosas funciones especficas. Los estudios de Ohno en la
dcada del 60 demostraron que el cromosoma X fue marcadamente conservado en tamao y
contenido de genes a lo largo de la evolucin de las especies euterianas. Las nicas excepciones son
los 3 genes localizados en el brazo corto del cromosoma X humano, que corresponden a genes
autosmicos en la especie murina. Son los denominados seudo autosmicos. El cromosoma Y es uno
de los menores cromosomas de la mayora de los mamferos euterianos, representa solamente el 2%
del complemento haploide y la porcin distal esta compuesta por secuencias repetidas y no
codificantes (Graves y Shetty, 2001). Adems, su tamao vara mucho en funcin de la variacin de
tamao del brazo largo. Alrededor de 70% del ADN del Y humano consiste de secuencias repetidas
del brazo largo. En un sentido evolutivo estas secuencias estn relacionadas con las secuencias del
cromosoma X (George y Wilson, 1994).
Estudios citogenticos y moleculares demostraron que el cromosoma Y es organizado por
diversas sub-regiones (Fig. 2). La regin del brazo corto posee una regin seudo autonmica
(autosomal) y la regin pericntrica que contiene el locus determinante del sexo. El brazo largo est
dividido en 2 regiones: la regin eucromtica que contiene genes involucrados en la
espermatognesis y la regin heterocromtica, que contiene genes la cual posee repetidas
secuencias de ADN y representa ms de la mitad del cromosoma Y (Cotinot y col.,1993; George y
Wilson, 1994). Se ha sugerido una posible accin del antgeno H-Y en los estadios precoces del
proceso de espermatognesis (McLaren, 1991).

Regin Telmero
Seudoautosmica

Genes determinantes del Brazo pequeo (Yp)


sexo

Centrmero

Regin eucromtica
Genes Brazo largo (Yq)
Espermatognesis

Regin heterocromtica Telmero

Fig 2. Modelo de cromosoma Y humano


Sexado 417
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