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The normal skin of the silkworm larvae is opaque and white in color and contains a large amount of uric acid, the endproduct of the nitrogen metabolism, in the hypodermis. In the " oily skin " mutants, however, the skin of the larvae is more or less translucent due to the reduced amount of uric acid contained in the hypodermis. It has been generally considered that the more transparent the skin becomes, the less uric acid it contains in the hypodermis. More than twenty such mutants are known so far and are located on the widely dispersed linkage groups, i. e., about half of the already identified twenty groups. This is in conspicuous contrast to the increasing evidences of existence of genic factors, with similar or related functions, occurring closely linked together on the chromosomes of the silkworm. Concerning the manifestation of genes, distribution of genes in the chromosomal set, and differentiation of chromosomes and genes, this group may provide a very fascinating problem to the genetics of the silkworm. From these points of view, we have been carrying out a genetical and biochemical analysis of the manifestation of this character on the basis of uric acid content in various genic combinations. Results of such an analysis are reported in this paper. The effects o f single and double doses of an autosomal oily gene, oa, on the uric acid contents were investigated by utilizing N1-deficiency which involves the oa locus. Results showed that the haplo-oa (oaf-) individuals contained more uric acid than the diplo-oa (oa/oa) ones (Tables 1 and 9). This suggests that oa gene is an antimorph in the sense that it inhibits the uric acid accumulation in the hypodermis. In the cas e of sex-linked genes, od and os, higher uric acid content was also obtained in the hemizygous individuals, suggesting the antimorphic nature of the genes (Tables 3 and 4). The normal alleles of oil y skin mutants are considered to be haplo-sufficient for the accumulation of uric acid since no or very few differences were found between single and double doses of the normal alleles (Tables 1, 2, 3, 4 and 5). The interactions of non-allelic oily genes on the uric acid accumulation were studied with the double oily mutants. Each of the oa (autosomal), os or od (sex-linked) genes was combined with one of the w3 allelic genes, w3, w01 and oew. It has been considered that the transparency in double mutants is determined predominantly by a gene having stronger effect on the transparency. This, however, does not hold true for all cases examined. In the combination of od with one of the w_3 a lleles, slight cumulative effect was observed toward reduction of uric acid content (Table 6) ; while in the case of oa combined with w_3 alleles much severe cumulative effect was observed, no or very few uric acid was detected in the double mutants (Table 9). In the case of os combined with w_3 alleles, however, higher uric a cid content as well as lower transparency was noted in the double mutants than in the single mutants, suggesting a quite distinct type of interactions (Tables 6, 7 and 8). In this case, a further interesting phenomenon was noted. Normally, uric acid contents in the w3 series mutants are in the order of w_3>w^M>oew. However, when each mutant is combined with os, it becomes equal in the double mutants, that is, os ; w_3=os; w^ol=os; oew. This result is interpreted to mean that all the w3 alleles must be dimorphic in nature of the locus ; one is a " common oily factor , " nonspecific to each alleles, and the other is a " specific oily factor, " peculiar to each mutant. This interpretation is consistent with the hypothesis of the complexity of this locus proposed by Chikushi previously . In the heterozygotes between w_3 alleles, a mutant having more translucent skin seems to be expressed as dominant (Tables 10 and 11). But careful observation suggests that this is not due to dominancy but to differences in transtype position effect caused by their inherent " specific oily factors." It is clear that the interaction of these genes is quite different depending on the combinations. From these results it is suspected that in spite of the phenotypic similarities, i. e., transparent skin and reduced uric acid accumulation in the hypodermis, the mode of action of each genes responsible for the oily skin characters is not al ways the same. Recently we have obtained some evide nces suggesting that individual genes controlling oily skin characters are primarily concerned with the alterations of the structure of the tissue proteins present in the hypodermis. It seems that the genic factors responsible for the production and differentiation of tissue proteins may be distributed on the various regions of different chromosomes. This may be the main reason why the genes in question are distributed so widely dispersed on the chromosomes of the silkworm.
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家蚕において油蚕性を発現する遺伝子として知られているものはすべて劣性でその種類は20数種に及んでいる.それらの属する連関群は既に判明しているだけでも第1,2,5,7,9,10,12,14,17,20染色体の多数に上るが,他に所属未詳のものも多いから,さらに広い分布を示すものであろうことが当然推察される.このように多数の,広汎に分布する異なつた遺伝子に支配される形質が程度の差こそあれ,皮膚の透明化,即ち油蚕性という単一の外的特徴を現わすのである.このような例は家蚕では他に見当らないのみでなく,広く動植物界を通じても極めて稀な現象であり,これに匹敵する事例としては数種の植物における広義の葉緑体異常(須藤,1955;高橋,1955;萩原,1955,等),Drosophila melanogasterにおけるヘテロクロマチン部の小欠失によるMinute形質の場合(Hannah,A.,1951)などを挙げうるに過ぎない.しかも家蚕において一般に類似した作用を現わす遺伝子は同染色体に集中分布する傾向がみられるのに比べると,油蚕性の場合は著しく対蹠的で,遺伝子の作用,分布及び分化の面から極めて興味深いものがある.この油蚕性に関する研究は本格的には田中義麿博士(1917,1921,1926,1929)の詳細な一連の遺伝学的研究を発端とする.しかして油蚕では皮膚細胞中に含有される尿酸量が少ないために皮膚が透明となるものであることがJucci(1932),畑村(1943),清水(1943)らによつて明らかにされて以来,油蚕は特に生理遺伝学的研究の好対象となり,数多くの研究が行なわれて来た.その結果,油蚕形質の多様性に関する生理学的観点からの多数の知見が得られるに至つたものの,主要観点においては正常(不透明性)に対する油蚕形質(透明性)という単相的な取扱いが行なわれて来たに過ぎないのである.また,その形質発現機構,すなわち皮膚細胞中の尿酸量を規制するという油蚕遺伝子の作用発現の面からの解析については,或は生成尿酸量自体の差と言い(畑村,1943;清水,1943;林,1961),或は皮膚細胞における尿酸吸着系の異常と言い(稲神・須藤,1955;江口,1961),未だ定説をみない現状である.ここにおいて著者らは油蚕性遺伝子の作用発現機構,油蚕遺伝子間の作用の異同を明らかにし,もつて家蚕における遺伝子分布傾向究明の一助とすべく,第1段階として皮膚尿酸量を標識とした油蚕性に関する遺伝子型による遺伝子作用効果の変動を調べることにした。まず伴性油蚕のos,od並びに常染色体のものとしでoaについて油蚕遺伝子のdosageeffectを調べ,遺伝子作用の性格,方向を明らかにし,ついで各々独立な油蚕遺伝子を組合せた二重突然変異系統を育成し,油蚕遺伝子間の相互作用を調べると共に,対立的に行動し,しかも透明度を異にするw_3系列油蚕についてホモ及び相互間のF_1での尿酸量を比較して遺伝子作用の異同を明らかにした.さらに湾系列油蚕の遺伝子構成についても分析した.かくて油蚕性の発現に関して新知見を加え得たと信ずるのでここにとりまとめて報告することにした.本文に入るに先立ち,終始懇篤なる助言と鞭達を賜つた林禎二郎教授に対し深甚の謝意を表する.
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