Bal arılarında yumurtanın yapısı ve embriyo gelişimi
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Date
2007
Authors
Akyol, Ethem
Journal Title
Journal ISSN
Volume Title
Publisher
Uludağ Üniversitesi
Abstract
Bu derleme en iyi gelişmiş sosyal böceklerden olan bal arılarında yumurtanın meydana gelişi, döllenmesi ve larval döneme kadar geçirdiği safhalar üzerine bilinenleri ve yakın zamanda yapılmış çalışmaları, taramak üzere hazırlanmıştır. Kraliçe arıların yumurtalıklarında bulunan yumurta tüplerinin uç kısımlarındaki doğurucu hücreler tarafından meydana getirilen yumurtalar yumurta tüpleri içerisindeki yaklaşık iki günlük yolculuk sırasında yine doğurucu hücreler tarafından meydana getirilen besleyici hücreler tarafından beslenerek gelişmelerini tamamlarlar. Gelişimini tamamlayan yumurta, yumurta tüplerinin dip kısmında besleyici hücrelerin kalan kısmını içine alır ve sonra da yumurta kabuğu oluşur. Yumurta daha sonra lateral oviductta ve median oviducta geçer. Median oviductan vajinaya geçen yumurta daha önce işçi arılar tarafından hazırlanmış ve kraliçe arı tarafından temizlik ve büyüklük bakımından kontrol edilmiş petek gözlerine bırakılır. Temiz olduğu anlaşılan ve çapı yaklaşık 5mm (işçi arı gözü) ve 8–9mm (kraliçe arı gözü) olan gözlere döllenmiş, 7mm (erkek arı gözü) olan gözlere ise döllenmemiş yumurtalar bırakılır. Yumurta vajinadan geçme esnasında spermatekadan gönderilen spermatazoa ile döllenirse dişi (kraliçe veya işçi arı) döllenmeden geçerse erkek bireyler meydana gelir. Petek gözlerine bırakılan yumurtalar ilk gün dik pozisyonda, ikinci gün eğik ve üçüncü gün ise göz tabanına yatık pozisyonda bulunurlar. Üç günlük bir süreden sonra açılarak önce larva halini alırlar ve bu süre larvanın cinsiyetine göre değişmek üzere 5 ila 7 gün arasında değişir. Bu sürenin sonunda ise göz kapanarak pupa dönemi başlar ve pupa dönemi de yine cinsiyete göre değişmek üzere 8 ila 14 gün arasında değişir. Pupa döneminin tamamlanmasından sonra ise göz açılır ve gelişmesini tamamlayan bireyler ergin olarak gözden çıkarak kolonideki görevlerini yerine getirmeye başlarlar.
This review aims to survey what is known and the relatively recent studies on formation and fertilization of the egg, and its development stages from embryo to larva in one of the most advanced social insects (Winston 1987, Moritz and Southwick 1992), the honey bee. The eggs are produced by the mother cells in the narrow end of the each tube of the ovaries. These eggs receive nutrients from the nurse cells during their development and travel in the egg tubules. Egg development takes approximately two days. The full-grown eggs absorb the nurse cell in the lower end of the tubules and the eggshell is formed. The mature eggs pass into the lateral oviduct and median oviduct. Then, they are discharged through the genital exit passage or vagina and are laid in the comb cells that were cleaned and prepared by the worker bees. Queens check cells before egg laying for cleaning and size. If eggs are fertilized while passing through the vagina they develop into female bees (queens or workers); those that are unfertilized become males or drones. After three days of embryonic development, these eggs laid in the comb cells turn into larvae that would develop into pupae and lastly, adult bees. Even though there are excellent reviews on the status quo, the new questions are not well represented in these reviews (e.g. Harbo and Rinderer, 1980, Koeniger 1986). Subjects where relatively recent information and questions are available follow: Gene expression during germ cell development is a question that could only recently be answered (e.g. Dearden et al. 2003, 2006). Similary, it is well known that only fertilized eggs develop into females, but the sex genes have been characterized only recently (see Beye et al. 2003). Since 1980s we know the importance of presence of multiple subfamilies in the honey bee colony (e.g. Page 1980), however, only recently do we see implication on worker reproduction. Worker reproduction is prevented by an elaborate mechanism including egg marker pheromones (e.g. Katzav-Gozansky et al. 2001, Martin et al. 2004) and worker policing (Oldroyd et al. 1991). Colony genetics and worker reproduction is of interest both to understand genetic variability in honey bee colonies and to examine impact of this variability on worker reproduction. Better assessment of genetic variation in colonies (e.g. Haberl and Moritz 1994) and even inside the spermatheca of the queen (e.g. Franck et al. 1999) have been the note-worthy improvements in this field. Another research frontier is to understand the order within chaos of queenless worker colonies (e.g. Neumann et al. 2000). Post-Embryonic development is the area where genetic and hormonal factors in development are being studied. The role of juvenile hormone in queen post-embryonic development has recently been re-visited for queens (Kaftanoglu et al. 2000). Cruz et al. (2002) examined changes in nuclear activity during post-embryonic development. Understanding egg production, fertilization, sex determination, and embryonic development could be important both for research on genetics and manipulation of the genome and for agricultural applications. Worker reproduction or laying workers, male production and queen production could be important for productivity, disease control by biological methods, and colony survival. This study compiled a survey of knowledge base and entry points to current literature on honey bee reproduction.
This review aims to survey what is known and the relatively recent studies on formation and fertilization of the egg, and its development stages from embryo to larva in one of the most advanced social insects (Winston 1987, Moritz and Southwick 1992), the honey bee. The eggs are produced by the mother cells in the narrow end of the each tube of the ovaries. These eggs receive nutrients from the nurse cells during their development and travel in the egg tubules. Egg development takes approximately two days. The full-grown eggs absorb the nurse cell in the lower end of the tubules and the eggshell is formed. The mature eggs pass into the lateral oviduct and median oviduct. Then, they are discharged through the genital exit passage or vagina and are laid in the comb cells that were cleaned and prepared by the worker bees. Queens check cells before egg laying for cleaning and size. If eggs are fertilized while passing through the vagina they develop into female bees (queens or workers); those that are unfertilized become males or drones. After three days of embryonic development, these eggs laid in the comb cells turn into larvae that would develop into pupae and lastly, adult bees. Even though there are excellent reviews on the status quo, the new questions are not well represented in these reviews (e.g. Harbo and Rinderer, 1980, Koeniger 1986). Subjects where relatively recent information and questions are available follow: Gene expression during germ cell development is a question that could only recently be answered (e.g. Dearden et al. 2003, 2006). Similary, it is well known that only fertilized eggs develop into females, but the sex genes have been characterized only recently (see Beye et al. 2003). Since 1980s we know the importance of presence of multiple subfamilies in the honey bee colony (e.g. Page 1980), however, only recently do we see implication on worker reproduction. Worker reproduction is prevented by an elaborate mechanism including egg marker pheromones (e.g. Katzav-Gozansky et al. 2001, Martin et al. 2004) and worker policing (Oldroyd et al. 1991). Colony genetics and worker reproduction is of interest both to understand genetic variability in honey bee colonies and to examine impact of this variability on worker reproduction. Better assessment of genetic variation in colonies (e.g. Haberl and Moritz 1994) and even inside the spermatheca of the queen (e.g. Franck et al. 1999) have been the note-worthy improvements in this field. Another research frontier is to understand the order within chaos of queenless worker colonies (e.g. Neumann et al. 2000). Post-Embryonic development is the area where genetic and hormonal factors in development are being studied. The role of juvenile hormone in queen post-embryonic development has recently been re-visited for queens (Kaftanoglu et al. 2000). Cruz et al. (2002) examined changes in nuclear activity during post-embryonic development. Understanding egg production, fertilization, sex determination, and embryonic development could be important both for research on genetics and manipulation of the genome and for agricultural applications. Worker reproduction or laying workers, male production and queen production could be important for productivity, disease control by biological methods, and colony survival. This study compiled a survey of knowledge base and entry points to current literature on honey bee reproduction.
Description
Keywords
Yumurta, Döllenme, Yumurtalık, Embriyo, Egg, Fertilization, Ovary, Embryo
Citation
Akyol, E. (2007). ''Bal arılarında yumurtanın yapısı ve embriyo gelişimi''. Uludağ Arıcılık Dergisi, 7(4), 135-144.