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BOR - Papers in Press, published online ahead of print April 28, 2004.
Biol Reprod 2004, 10.1095/biolreprod.104.028282
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Submitted February 6, 2004
Returned for revision February 24, 2004
Accepted April 19, 2004

Gamete Biology


Intraoocyte Localization of MAD2 and Its Relationship with Kinetochores, Microtubules, and Chromosomes in Rat Oocytes During Meiosis

Dong Zhang , Wei Ma , Yong-Hai Li , Yi Hou , Shi-Wen Li , Xiao-Qian Meng , Xiao-Fang Sun , Qing-Yuan Sun , and Wei-Hua Wang *

* To whom correspondence should be addressed. E-mail: wangweihua11{at}yahoo.com.

Abstract
The present study was designed to investigate subcellular localization of MAD2 in rat oocytes during meiotic maturation and it's relationship with kinetochores, chromosomes and microtubules. Oocytes at germinal vesicle (GV), prometaphase I (ProM-I), metaphase I (M-I), anaphase I (A-I), telophase I (T-I) and metaphase II (M-II) were fixed and immunostained for MAD2, kinetochores, microtubules and chromosomes. The stained oocytes were examined by confocal microscopy. Some oocytes from GV to M-II stages were treated by a microtubule disassembly drug, nocodazole or treated by a microtubule stabilizer, Taxol before examination. Anti-MAD2 antibody was also injected into the oocytes at GV stage and the injected oocytes were cultured for 6 h for examination of chromosome alignment and spindle formation. It was found that MAD2 was at the kinetochores in the oocytes at GV and ProM-I stages. Once the oocytes reached M-I stage in which an intact spindle was formed and all chromosomes were aligned at the equator of spindle, MAD2 disappeared. However, when oocytes from GV to M-II stages were treated by nocodazole, spindles were destroyed and MAD2 was observed in all treated oocytes. When nocodazole-treated oocytes at M-I and M-II stages were washed and cultured for spindle recovery, it was found that once the relationship between microtubules and chromosomes was established, MAD2 disappeared in the oocytes even though some chromosomes were not aligned at the equator of spindle. On the other hand, when oocytes were treated with Taxol, MAD2 localization was not changed and was the same as that in control. However, immunoblotting of MAD2 indicated that MAD2 was present in the oocytes at all stages; nocodazole and Taxol treatment did not influence the quantity of MAD2 in the cytoplasm. Significantly higher proportions of anti-MAD2 antibody injected oocytes proceeded to premature A-I stage and more oocytes had misaligned chromosomes in the spindles. The present study indicates that MAD2 is a spindle checkpoint protein in rat oocytes during meiosis. When the spindle was destroyed by nocodazole, MAD2 was re-activated in the oocytes to overlook the attachment between chromosomes and microtubules. However, in this case, MAD2 could not check unaligned chromosomes in the recovered spindles suggesting that a normal chromosome alignment is maintained only in the oocytes without any microtubule damages during maturation.

Key words: Gamete Biology • Gametogenesis • Meiosis • Oocyte development


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