What the books sayAntonio’s ImpactDevelopmental Genetics of Drosophila. A. Ghysen (Ed).Bilbao: UBC Press. International Journal of DevelopmentalBiology, vol. 42, no. 3, pp 229–5401

Reviewed byAdam S. WilkinsBioEssays Editorial OfficeCambridge, UK

Antonio Garcia-Bellido (AGB) is undeniably one of the semi-nal figures of 20th century developmental biology. His signifi-cant contributions have been numerous but, perhaps, themost important were the concepts of compartments andselector genes, which were first elaborated in a talk in 1974 atthe then Ciba Foundation (now the Novartis Foundation).These ideas, which were derived from a powerful methodol-ogy and an intriguing set of results, revolutionized thinking indevelopmental genetics and were a powerful stimulus to itsgrowth. In addition to his many direct contributions, however,AGB has had an enormous influence through the legions ofyoung investigators whom he has trained and/or inspired. Intribute to his many gifts to the field, a Festschrift, under theeditorship of Alain Ghysen, was put together and publishedlast year in the International Journal of Developmental Biol-ogy.1 Altogether this volume consists of 37 articles, twointerviews and seven memoir-reminiscences.

One of the pleasures of reading this collection of articles isits multi-level diversity. One element of the diversity consistsof the contributors themselves, who range from major andsenior figures, including undeniable Grand Old Men of biol-ogy (including Francois Jacob, Ed Lewis, Lewis Wolpert, andGunther Stent) to many younger scientists at the postdoctorallevel. Though the latter inevitably lack some of the experienceand broader perspective of the older scientists, some of theircontributions are excellent, amongst the best in the book.Another aspect of the diversity is in the range of the subjectmatter. While most of the articles are about aspects of thedevelopmental biology of Drosophila, with only a handfuldevoted to vertebrate systems, the specific topics range fromearly egg and embryo development to the structure of geneticnetworks to neural development and behavior.

A third kind of diversity is in the style of treatments, whichis not surprising in a collection featuring so many authors.Several articles are straightforward reports, emphasizing thework of the authors’ labs. Most, however, are broader reviewsof subjects, usually introduced with a few words about AGB’sinfluence on the work. A few articles, such as John Edwards’tribute to Vincent Wigglesworth and Rene Thomas’ lucidexposition of the role of positive feedback circuits in drivingdevelopmental decisions and Jonathan Hodgkins’ dissectionof seven different forms of mutational pleiotropy and whatthey can reveal are, at most, only tangentially connected toAGB’s body of work. Yet they are no less enjoyable, informa-tive or interesting for that. One suspects that the editorencouraged and actively sought this diversity of subject andapproach.

There is also one piece of comedy. Lewis Wolpert hadbeen unable to contribute an article, hence the editor did ajoint interview with him and AGB. Both Lewis and AGB arerenowned as brilliant talkers. On the other hand, should thereever be a Hall of Fame of Great Listeners, it is unlikely thateither would even be nominated for it, let alone elected. Notsurprisingly, the interview shows two brilliant men largelytalking past one another. (The image of AGB as a relentlessyet always fascinating and informative talker comes through,loud and clear, you might say, in many of the reminiscences.)Yet, this exchange between Lewis and AGB is fun to read. Italso has a serious, if insufficiently-explored difference in pointof view. Lewis tends to emphasize the reactions of cells, notleast their mechanical reactivities, to ‘‘information’’ suppliedfrom outside while Antonio emphasises the ‘‘personality’’ (hisword) of individual cells, and their consequent roles as actorsin the drama of development, as determined by their ex-pressed genes. Clearly, both elements are important indevelopment and both men pay lip service to the other’sconcerns. Nevertheless, developmental biology still awaits aconvincing synthesis of these two perspectives.

Despite the range of subject matter, three subjects are thefocus of many of the contributions, and they are closelyassociated with AGB’s work: the mechanisms of spatialpatterning in general, the roles of Hox and other ‘‘selectorgenes’’ more specifically, and the control of cell proliferationas an element of patterning during development. Several ofthe exciting papers on pattern formation are: the contributionfrom Mar Ruiz-Gomez on muscle patterning, discussions onthe elements of patterning in the wing by Alicia Hidalgo andJose de Celis, and a review by Juan Pablo Couso and SarahBishop on the patterning of the fruit fly leg along theproximo-distal axis.

With respect to the roles of Hox genes, the contributionsfrom James Castelli-Gair and Michael Akam explore anambiguity in AGB’s original depiction of selector genes. Thisconcerns the question: does each selector gene convey ablanket area-identity (for compartments or segments) or does

1Though published as an issue of the IJDB, this volume can be boughtseparately from the publishers. The price is 7000 pesetas, whichincludes handling and airmail postage, and is payable by Visa/Mastercard or check. Payment by credit card can be by letter, fax orE-mail with card details and should be sent to: University of the BasqueCountry Press/Journals Subscription Department, Post Box 1397,E-48080 Bilbao, Spain. Tel: (34)-(94)-601 2000 Ext. 5126; Fax:(34)-(94)-480 1314; E-mail: luxedito@lg.ehu.es

What the books say

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it control individual cell/organ properties within that area, thearea property being the additive effect of those individualproperties? Both of these articles argue for the latter, thoughin slightly different ways. Mike Akam, in particular, describesthe roles of enhancers as integrators and fine-tuning agentsof developmental, and evolutionary, change—a view that onealso associates with Eric Davidson. The importance of dis-crete enhancer elements in gene regulation is similarly one ofthe themes of an excellent review of the Achaete-Scutecomplex by Juan Modolell and Sonsoles Campuzano.

With respect to the control of cell proliferation, there areseveral important contributions by, amongst others, AGBhimself and his son, Antonio. The two Antonio’s describe their‘‘Entelechia’’ hypothesis on how growth is organized so as toproduce the correct sizes and proportions of pattern elementswithin compartments. It is hard not to wonder: how testablewill this idea prove to be? Only the future will tell but the ideais, in AGB’s best tradition, incisive and stimulating. A ratherdifferent, but equally provocative and interesting view, of cellproliferation control (at compartment boundaries) is providedby David Gubb, who also provides an interpretation of howcell polarity properties shape patterning.

There are too many good articles in this volume to list themall in a short review. Furthermore, each reader will have hisown list of favorites. Since a reviewer who likes a book,however, should also register a few quibbles—in order toprove that he was reading it attentively—I will mention two.First, there is practically no mention of the contributions of

Curt Stern, yet it was Stern who developed the technique ofstudying mitotic recombination that played so major a part inAGB’s development of the concept of compartments. Onemight even say that it was part of AGB’s genius to haveapplied Stern’s technique (mitotic recombination) to Ed Lewis’problem (homeotic control genes). Gunther Stent’s history ofcell lineage studies really should have mentioned Stern’scontribution to the development of developmental genetics.Second, I would have been curious to have AGB’s ownperspective on his 1974 ideas. The implications of his fewremarks in the interview at the front of the volume suggestthat his views are unchanged. Can this really be the case ordid the interview format somehow crimp his explanation? Weknow, for instance, that the genes immediately downstreamof selector genes are not just ‘‘realizator genes’’, that is celldifferentiation functions in the original formulation, but includemany regulator genes, as is described in the article byJacques Pradel and Robert White.

Altogether, the editor, the authors, and the publishers havedone a fine job. The articles are liberally illustrated with goodfigures, the number of typos is hardly zero but is not too high,and the European-flavour of the English of many of thecontributors is delightful and contributes to the ‘‘personality’’of the articles. Though, inevitably, some of the articles will beovertaken by new findings and interpretations, the majorityare sufficiently broad and thoughtful to be valuable for manyyears to come. One hopes that Antonio was pleased with theresult.

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