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Heteroblasty—A Review

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Abstract

Virtually all plants show a certain degree of variation among individual metamers during ontogeny. In some cases, however, there are abrupt and substantial changes in form and function (e.g. in leaf form, leaf size, phyllotaxy, internode length, anthocyanin pigmentation, rooting ability, or wood structure). These plants were called “heteroblastic” by Karl Goebel more than a century ago, but the functional significance of this type of ontogenetic change, the evolutionary trajectories in different plant groups, even their frequency in the plant kingdom are still unresolved issues. We argue that slow progress is partly due to an on-going terminological confusion and the lack of distinction between other developmental processes such as ontogenetic drift. This review develops a conceptual framework for future scientific work, proposes a quantitative index of heteroblasty, and discusses the evidence for developmental regulation, functional significance, and evolutionary implications of heteroblasty to provide a stimulating basis for further research with this fascinating group of plants.

Zusammenfassung

Fast alle Pflanzen zeigen während der Individualentwicklung ein gewisses Maß an Variabilität einzelner Metamere. Bei manchen Arten kommt es jedoch zu einer ausgeprägten und sprunghaften Änderung in Form und Funktion (z.B. in Blattform oder -stellung, Internodienlänge, Pigmentierung, Holzstruktur, oder hinsichtlich der Fähigkeit zur Bildung von Adventivwurzeln). Obwohl diese Arten von Karl Goebel schon vor mehr als einem Jahrhundert als “heteroblastisch” beschrieben wurden, sind die funktionelle Bedeutung des Phänomens, dessen Evolution innerhalb einzelner Pflanzengruppen, wie auch die Häufigkeit im Pflanzenreich immer noch weitgehend ungeklärt. Dieser schleppende Fortschritt mag mit einem weit verbreiteten terminologischen Durcheinander und dem Fehlen einer klaren Abgrenzung von anderen Entwicklungsprozessen wie der “ontogenetischen Drift” zusammenhängen. Unser Übersichtsartikel entwickelt einen klaren konzeptionellen Rahmen, um eine Basis zu schaffen für zukünftige Forschungsarbeiten dieses faszinierenden Entwicklungsphänomens. Dazu schlagen wir einen quantitativen Index der Heteroblastie vor, skizzieren den gegenwärtigen Wissensstand der Regulierung von Entwicklungsprozessen bei Pflanzen, die bisherigen Untersuchungen zur funktionelle Bedeutung heteroblastischer Veränderungen, beleuchten aber auch die evolutionären Implikationen der Heteroblastie.

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Acknowledgements

Of tremendous help in procuring the literature for this review were Herta Sauerbrey (Oldenburg, Germany) and Angel Aguirre (STRI, Panama). Financial support of the Deutsche Forschungsgemeinschaft (GZ ZO 94/4-1) for our current studies on heteroblasty in bromeliads is acknowledged. The Deutsche Forschungsgemeinschaft also funds ongoing research in AB’s laboratory (BE 2547/7-2 and/8-1). Critical comments on an earlier draft by Dirk Albach, Oldenburg, are appreciated.

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Appendix I

Appendix I

A diverse body of literature that deals with heteroblasty has accumulated over the last 100 or so years. Unfortunately, the older literature is hardly covered in data bases such as WOS, in part because quite a few contributions were made in monographs, in dissertations, or in journals, which are not indexed. Other studies are not readily accessible due to terminological confusion. To assist in future research we have done an extensive literature search and compiled the following list of research articles, books and book chapters that deal with heteroblasty. Also included is a selection of general textbooks, which provide different definitions of “heteroblasty”, “heterophylly” and/or “phase change”.

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Zotz, G., Wilhelm, K. & Becker, A. Heteroblasty—A Review. Bot. Rev. 77, 109–151 (2011). https://doi.org/10.1007/s12229-010-9062-8

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