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絶対単為生殖種キイロヒメアリの繁殖生態と社会構造

井戸川, 直人 京都大学 DOI:10.14989/doctor.k24300

2022.11.24

概要

多くの動物は雌雄の配偶子を接合させる有性生殖により繁殖している。増殖に直接貢献しないオスの生産はコストであるにもかかわらず、メスのみで繁殖する単為生殖を行なう種は少ない。また、とりわけ社会性昆虫にとっては、単為生殖は家族集団の血縁度を高めるため、包括適応度の観点から社会性昆虫に高い利益をもたらしうる。しかし、代表的な社会性昆虫であるアリ類において、オスをともなわない絶対産雌性単為生殖が報告されている種は全体の0.1%に満たない。このような種の生活史を明らかにすることは、社会性昆虫の繁殖システムの多様性と進化の理解に寄与する。また、単為生殖の結果として生じる親子間やコロニー内の特殊な遺伝構造は、真社会性の基盤である分業システムや発生の可塑性のユニークなモデル系として利用できる。本研究では、オスが発見されておらず、女王が絶対産雌性単為生殖によりワーカーと次世代の女王を生産すると考えられるキイロヒメアリMonomorium trivialeに着目し、その生活史を包括的に解明することを目的とした。本論文は以下のように要約される。

 第1章では、単為生殖性の生物を対象とした研究の意義について述べるとともに、これまでに社会性昆虫から報告されてきた単為生殖を組み込んだ繁殖システムの類型を概観した。さらに単為生殖性の社会性昆虫が有望な研究モデルであることを示したうえで、新規モデルとしてキイロヒメアリに着目する理由を述べた。

 第2章では、キイロヒメアリの単為生殖システムを多角的に検証した。国内の分布域をカバーする全国各地の個体群を用いて、未交尾女王の飼育実験とマイクロサテライト解析の結果、すべての調査個体群が絶対産雌性単為生殖を行っていることが明らかになった。また、国内の各個体群のミトコンドリアDNA配列に基づく年代推定を行なったところ、個体群間の変異は小さく、2~7万年以内に生じうる程度のものであると推定された。このことから、本種の単為生殖系統が国内において急速に分布を拡大したことが示唆された。さらに産雌性単為生殖の起源に着目し、キイロヒメアリの体内細菌叢を解析したところ、膜翅目において単為生殖を誘導する細胞内共生細菌は検出されず、本種の産雌性単為生殖能力は内因性であると考えられた。なお解剖学的観察の結果、ワーカーは卵巣を喪失しており、女王による絶対産雌性単為生殖はワーカーが完全不妊の種からのみ進化するという先行研究の見解と一致した。

 第3章では、キイロヒメアリのカースト発生に着目し、女王とワーカーの発生段階を記載した。形態計測と飼育実験の結果、将来女王となる終齢幼虫は特異的な突起状の体表構造をもつことが分かった。このような女王特異的な構造はこれまで報告されておらず、アリ類の幼虫における新奇カースト二型として位置付けられた。さらに発生過程の観察からは、女王個体の蛹化時に機能的な翅が形成されないことも明らかとなった。このことから、本種の女王は飛行能力をもたず、歩行によって分散し新巣を創設すると考えられた。長距離分散できない無翅女王の発見は、絶対産雌性単為生殖を行なうアリにおいて初めてであり、アリ類における個体レベルと巣レベルの繁殖システムの新たな組み合わせを見出した。

 第4章では、絶対産雌性単為生殖という個体レベルの繁殖システムと、無翅女王による短距離分散という巣レベルの繁殖システムの組み合わせがもたらす空間遺伝構造に着目した。京都府内のキイロヒメアリ集団についてRAD-seq法を用いてSNP多型を検出し、集団遺伝学的解析を行なったところ、異なる採集サイト間で遺伝的分化が認められた。これは、本種が長距離分散できないことと整合する結果と考えられた。一方で、各採集サイト内における巣間の遺伝的分化は小さく、それぞれの巣には遺伝的に均一でない個体が含まれていた。さらに行動観察により、本種のワーカーは異なる巣の同種個体にも敵対行動を示さないことも確かめられた。一連の結果は、離れた巣の個体同士の直接的な相互作用が不可能なほどの地理的スケールにおいて、個体群を単一のコロニーとみなすことができるという融合コロニー性の定義に当てはまる。これによりキイロヒメアリは、絶対産雌性単為生殖を行なうアリにおいて、融合コロニー性が認められた初めての例と位置付けられた。女王アリが絶対産雌性単為生殖を行ない、かつ長距離分散できない本種では、隣り合う巣の遺伝的距離がきわめて近くなり、巣仲間識別の標識となるアレル(対立遺伝子)の多様性が失われた結果、この特殊な社会構造が獲得されたと考えられる。

 第5章では本研究の主要な成果を要約するとともに、キイロヒメアリと既知の侵略的外来アリとの生活史形質の共通点を挙げて、自然分布域外に移入された場合の潜在的リスクを指摘した。また、産雌性単為生殖を行なうアリを用いた研究の今後の展望を述べた。

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