Sexado 415

SELECCIÓN DEL SEXO EN MAMÍFEROS

Ligia M Cantarelli Pegoraro

Maria Gabriela T Rheingantz
& Gustavo A Palma

Introducción

El origen de la determinación sexual de la descendencia de los mamíferos siempre fue motivo

de especulaciones. Hace más de 3000 años, varias teorías, suposiciones y creencias fueron
postuladas. En la Grecia antigua los filósofos meditaban sobre la teoría de la determinación sexual,
según el lado izquierdo o derecho y del estado caliente o frío. Se especulaba que el testículo del lado
derecho originaba descendencia macho y el lado izquierdo hembras. El lado del útero materno fue
considerado también un factor determinante. La posición durante el acto sexual (norte/sur) favorecía
la generación de machos y hembras. Los estudios científicos que colaboraron con la dilucidación de
los factores involucrados en la determinación sexual comenzaron efectivamente con el
descubrimiento del espermatozoide, en el siglo XVII, su origen y función en el año 1841, con el
descubrimiento de la gameta femenina en el año 1827 y con la constatación de la ocurrencia de la
fecundación, solamente al final del mismo siglo. El misterio de la generación fue dilucidado al final del
siglo XIX con el descubrimiento de los cromosomas. En esta época se consideraba también que las
condiciones ambientales eran determinantes para el sexo de los productos, que las condiciones
nutricionales adversas originaban machos, mientras que las favorables hembras. Así 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 mamíferos se estableció la importancia de los cromosomas sexuales X e Y,
siendo el Y responsable por la generación de productos del sexo masculino (ver revisión sobre
mitología de la determinación del sexo de Beatty, 1974, 1978; Gledhill, 1988a, b; Mittwoch, 2005).
En mamíferos euterianos se determinó que la diferenciación sexual es un proceso secuencial
que se inicia con el establecimiento del sexo genético o cromosómico (cromosomal) en el momento
de la fecundación (XX o XY). El espermatozoide portador del cromosoma Y determinará la formación
de un individuo del sexo masculino y el espermatozoide portador de X uno de sexo femenino. La
primera fase de la diferenciación sexual es denominada determinación primaria del sexo y es cuando
tiene lugar la organogénesis gonadal, en función del sexo genético del embrión, culminando con la
formación del fenotipo sexual. Cada etapa de este proceso es dependiente de la etapa precedente y,
en circunstancias normales, el sexo cromosómico está de acuerdo con el sexo fenotípico (Jost y col.
1973; George y Wilson, 1994).

Sexo Sexo Sexo

Cromosómico Gonadal Fenotípico

Fig 1. Esquema de la determinación sexual en los mamíferos

Tanto el macho (XY) como la hembra (XX) poseen gónadas idénticas en la fase inicial del
desarrollo, cuando la cresta gonadal es denominada gónada indiferenciada o gónada totipotente. La
segunda fase de la diferenciación sexual abarca el desarrollo de la gónada indiferenciada en
testículos y ovarios. La tercera fase es la diferenciación de los conductos que consiste en la formación
de los conductos de Wolf en el embrión macho y de los conductos de Müller en embrión de sexo
femenino. Dos hormonas secretadas por el testículo, antimülleriana (AMH) y testosterona,
transforman entonces el tracto urogenital indiferenciado en tracto genital característico del macho. La
hormona antimülleriana es secretada por las células de Sertoli y causa regresión de los conductos
416 Pegoraro, Rheingantz & Palma
femeninos (de Müller) en el embrión masculino. La testosterona, secretada por las células de Leydig
fetales es responsable del restante desarrollo del tracto reproductivo masculino, incluido la
estabilización y diferenciación de los conductos de Wolf en los órganos secundarios masculinos y la
formación de los genitales externos y próstata. La última fase es la diferenciación del sinus urogenital
y de los genitales externos, de acuerdo con el sexo del producto (Josso y col. 1977; Baskur, 1980).
En los mamíferos euterianos, el cromosoma X es grande, representa 5% del genoma haploide y
es portador de genes que codifican numerosas funciones específicas. Los estudios de Ohno en la
década del 60 demostraron que el cromosoma X fue marcadamente conservado en tamaño y
contenido de genes a lo largo de la evolución de las especies euterianas. Las únicas excepciones son
los 3 genes localizados en el brazo corto del cromosoma X humano, que corresponden a genes
autosómicos en la especie murina. Son los denominados seudo autosómicos. El cromosoma Y es uno
de los menores cromosomas de la mayoría de los mamíferos euterianos, representa solamente el 2%
del complemento haploide y la porción distal esta compuesta por secuencias repetidas y no
codificantes (Graves y Shetty, 2001). Además, su tamaño varía mucho en función de la variación de
tamaño del brazo largo. Alrededor de 70% del ADN del Y humano consiste de secuencias repetidas
del brazo largo. En un sentido evolutivo estas secuencias están relacionadas con las secuencias del
cromosoma X (George y Wilson, 1994).
Estudios citogenéticos y moleculares demostraron que el cromosoma Y es organizado por
diversas sub-regiones (Fig. 2). La región del brazo corto posee una región seudo autonómica
(autosomal) y la región pericéntrica que contiene el locus determinante del sexo. El brazo largo está
dividido en 2 regiones: la región eucromática que contiene genes involucrados en la
espermatogénesis y la región heterocromática, que contiene genes la cual posee repetidas
secuencias de ADN y representa más de la mitad del cromosoma Y (Cotinot y col.,1993; George y
Wilson, 1994). Se ha sugerido una posible acción del antígeno H-Y en los estadios precoces del
proceso de espermatogénesis (McLaren, 1991).

Región Telómero
Seudoautosómica

Genes determinantes del Brazo pequeño (Yp)

sexo

Centrómero

Región eucromática
Genes Brazo largo (Yq)
Espermatogénesis

Región heterocromática Telómero

Fig 2. Modelo de cromosoma Y humano

Sexado 417
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