Facing the future of plant-insect interaction research: Le Retour a la "Raison d'Etre''
Facing the future of plant-insect interaction research: Le Retour a la "Raison d'Etre''
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DOI：
10.1104/pp.107.113472
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发表时间：
2008-03-01
期刊：
影响因子：
7.4
通讯作者：
Zangerl, Arthur R.
中科院分区：
文献类型：
--
作者：
Berenbaum, May R.;Zangerl, Arthur R.
草食性昆虫和它们所消费的被子植物之间的相互作用，共同构成了陆地群落中的大多数宏观物种，经常被比喻为战争（例如Gonzalez和Nebert，1990），而往复防御和反防御的过程被称为共同进化的军备竞赛（Whittaker和Feeny，1971）。然而，可能很少有人意识到，这一特殊的研究领域是作为一场实际的而不是隐喻的战争的直接后果而兴起的。当第二次世界大战爆发时，伦敦帝国理工学院的动物学和应用昆虫学系搬迁到斯劳，并成立了害虫感染实验室来帮助战争。昆虫生理学家戈特弗里德·弗伦克尔（Gottfried Fraenkel）对储藏产品害虫的营养进行了研究，以深入了解如何控制它们。在这个过程中，他确定，除了少数例外，人类和昆虫的营养需求基本上是相同的，而且，大多数绿色植物在营养上基本相等（Fraenkel，1953）。这一观察结果反过来又导致了这样的猜测，即无营养物质，即所谓的次生物质，特异地分布在整个植物界，决定了寄主植物的利用模式--“次生植物物质的分布和组成的巨大变化，当时还没有全面和合理的解释，解释了同样惊人的昆虫-食物-植物关系的变化，通过它们作为昆虫和其他生物的驱避剂和引诱剂“（Fraenkel，1984，第1页）。1953年在哥本哈根举行的一次国际动物学大会上，弗兰克尔首次提出了革命性的新观点，但直到1959年，他才在《科学》杂志上发表了一篇名为《次生植物物质存在的理由》的文章。植物产生如此多样的次级代谢物是为了抵御昆虫和其他食草动物，这种观点实际上是缓慢流行的;这篇文章在1959年至1964年期间被引用不到12次。然而，埃利希和瑞文（1964）的文章重新引起了人们的注意，他们通过对蝴蝶和植物的系统发育分析扩展了这一观点，并提出共同进化（这个过程弗伦克尔早先称之为“相互适应辐射”）不仅是植物次生物质巨大多样化的原因，也是被子植物和昆虫多样化的原因。从技术上讲，2003年是植物和昆虫之间化学共同进化概念的50周年纪念日。由于这个想法直到1964年才获得太多的关注，也许2014年是更合适的50周年纪念日;然而，2009年标志着化学共同进化概念在主流文献中的50周年。因此，随着这一年的临近，现在似乎应该评估特别是近年来取得的进展，并确定仍然存在的挑战。1959年，为了证明自己的观点，弗伦克尔将他的论点集中在少数几个植物科上，这些科的信息与它们的次生化学以及它们与食草昆虫的生态联系有关。其中包括十字花科、伞形科、豆科、茄科、桑科和禾本科（尽管十字花科、伞形科、豆科和禾本科，由于分类学的一致性，现在分别被称为禾本科、伞形科、豆科和禾本科）。
The interaction between herbivorous insects and the angiosperm plants they consume, collectively constituting the majority of macroscopic species in terrestrial communities, has often been metaphorically likened to warfare (eg Gonzalez and Nebert, 1990), and the process of reciprocating defense and counter defense has been called a coevolutionary arms race (Whittaker and Feeny, 1971). Few people probably realize, however, that this particular field of study arose as a direct consequence of an actual, rather than metaphorical, war. When World War II broke out, the Department of Zoology and Applied Entomology at Imperial College, London, was relocated to Slough and the Pest Infestation Laboratory was founded to aid in the war effort. Insect physiologist Gottfried Fraenkel undertook a study of the nutrition of stored product pests in an effort to gain insight into how to control them. In the process, he determined that, with only a few exceptions, human and insect nutritional requirements are essentially the same and that, moreover, the majority of green plants are essentially nutritionally equivalent (Fraenkel, 1953). This observation in turn led to speculation that nonnutritious substances, the so-called secondary substances idiosyncratically distributed throughout the plant kingdom, determine patterns of host plant utilization—‘‘the enormous variety in the distribution and composition of the secondary plant substances, for which no comprehensive and plausible explanation then existed, accounted for the equally staggering variety of insect-food-plant relationships, by their acting as repellents and attractants for insects and other organisms’’(Fraenkel, 1984, p. 1). Fraenkel’s revolutionary new idea was first advanced in a lecture at an international zoological congress in 1953 held in Copenhagen, but it did not reach the mainstream press until 1959, when he authored a now-famous article titled ‘‘The raison d’être of secondary plant substances’’in the journal Science. The idea that the reason plants manufacture such a diversity of secondary metabolites is to defend themselves against insects and other herbivores was in fact slow to catch on; the article was cited fewer than 12 times between 1959 and 1964. Attention was refocused on the article, however, by Ehrlich and Raven (1964), who expanded on the idea with a phylogenetic analysis of butterflies and plants and suggested that coevolution (the process Fraenkel had earlier called ‘‘reciprocal adaptive radiation’’) was responsible not only for the tremendous diversification of plant secondary substances, but also for the diversification of angiosperm plants and the insects that eat them. Technically, then, the 50th anniversary of the concept of chemical coevolution between plants and insects passed without notice in 2003. Inasmuch as the idea did not gain much traction until 1964, perhaps a more appropriate 50th anniversary would be 2014; yet 2009 marks 50 years of the concept of chemical coevolution in the refereed mainstream literature. Thus, as the year approaches it seems timely to evaluate the progress that has been made, particularly in recent years, and to identify the challenges that remain. To make his case in 1959, Fraenkel concentrated his argument on a handful of plant families for which information was available relating to their secondary chemistry and to their ecological associations with insect herbivores. These included the Cruciferae, Umbelliferae, Leguminosae, Solanaceae, Moraceae, and Gramineae (although Cruciferae, Umbelliferae, Leguminosae, and Gramineae, for reasons of taxonomic consistency, are now known as Brassicaceae, Apiaceae, Fabaceae, and Poaceae, respectively …