Mushrooms follow an almost embryo-like developmental pathway. These are microscope images of sections cut right through the centres of developing fruit body 'initials' of the field mushroom Coprinus cinereus; the youngest is on the left of the panel.

You can see that a 'young mushroom' is clearly established well before the initial reaches one millimetre high. The mature mushroom will be 100 mm tall, so this is a tiny embryo by comparison.

Homologues and analogues of all of the developmental mechanisms known in animals and plants can be found
in fungi
mechanical effects
temporal sequencing
pattern formation and morphogenetic fields
reaction with extracellular matrix
a fungal kind of programmed cell death

But there are differences. For example, hyphal cells require continuous reinforcement of their differentiation 'instructions'. This reinforcement is part of the context within which they normally develop and if they are removed from it then most differentiated fungal cells can revert to being undifferentiated hyphae.

Fungi are modular organisms

Remember that mushrooms not individual organisms. Mushrooms are appendages (literally 'fruits') of their mycelium. In fact, fungi are 'modular organisms', like clonal corals and vegetatively-propagated plants.

The nature of modular organisms is discussed in these two references:

Andrews, J. H. (1995). Fungi and the evolution of growth form.
Canadian Journal of Botany 73, S1206-S1212.
Harper, J. L. et al. (1986). The Growth and Form of Modular Organisms.
The Royal Society: London.

Developmental subroutines

Fungal morphogenesis is organized into a collection of distinct developmental processes, called 'subroutines'.
These are recognizable at all levels, and are genetically and physiologically distinct. Developmental subroutines
may run in parallel or in sequence. Normal morphogenesis is made up of a range of developmental subroutines:
subroutines for hymenophore, hymenium, stem, cap, gills, veil, etc.
subroutines can be put into operation independently of one another
under separate genetic control
under separate physiological control

When played out in their correct arrangement, ‘normal’ morphology is obtained. Subroutines may be disabled genetically or through physiological stress. If some subroutines are disabled, an abnormal morphology results.

Twin fruit bodies of Clitocybe nebularis Triple fruit bodies of Melanoleuca melaleuca Twin fruit bodies of Clitocybe nebularis
Photos by H. van der Aa, from Mycologist, volume 11, part 2 (May 1997), p. 81
This close up shows how one fruit body of Clitocybe nebularis has burst out of the internal tissues of its twin. For some reason, the upper fruit body formed in a completely inappropriate position - within the cap of a pre-existing fruit body. In these examples, the subroutines for fruit body initiation and development were put into effect in the wrong place and at the wrong time, but they otherwise gave rise to an essentially normal morphology. 
Other examples show that 'fruit bodies' very different from the normal morphology can be produced.

 

This is what the fruit bodies of Volvariella bombycina normally look like ... 

and the pictures below show some abnormal ones; from left - a Morchella-like fruit, an inverted cap and a club-shaped fruit body.

 

Developmental errors are not limited to fungi.

10 to 15% of human embryos suffer major malformations (defined as primary errors of development) soon after conception - the majority are aborted. About 0.7% of newborns have multiple major malformations, about 3% have one major malformation and 14% have a single minor malformation.

The difference with fungi, is that most of the abnormal fungal fruit bodies still produce spores. That is, they are still functional. It seems that fruit body abnormalities in fungi demonstrate their flexibility in expression of their developmental subroutines in ways that allow the fruit body to react to adverse conditions and still produce a crop of spores.

I call this tolerance of imprecision. Fungal fruit bodies can tolerate imprecision of their development, and this ability to tolerate uncertainty in development is what makes the fungi so successful.

For the full story ...

Fungal Morphogenesis

by David Moore

 

published by Cambridge University Press, New York Office

(xiv + 469 pages; ISBN 0 521 55295 8)

YOU CAN ORDER the book from the publisher, Cambridge University Press, at this URL: http://www.cup.cam.ac.uk/Scripts/webbook.asp?isbn=0521552958

The book brings together for the first time the full scope of fungal developmental biology, providing a coherent account which will be the basis for research in the future. The treatment also releases fungal morphogenesis from the confines of mycology, showing how and why this eukaryotic Kingdom deserves to be in the mainstream of developmental research. The author's view is quite simply that if you are ignorant of fungal morphogenesis then your understanding of developmental biology is incomplete.

The book is aimed at all biologists. Throughout, the author blends together physiological, biochemical, structural and molecular descriptions within an evolutionary framework, combining the older literature with the most recent. A comprehensive description of fungi is not attempted, though sufficient information is provided about fungal biology to give the general reader a rounded view of the mycological context within which fungal morphogenesis is played out, without obscuring the broader biological significance. Jargon is avoided, technical terms demystified and a reader with knowledge of basic biology should not need to bring any other knowledge with him/her, nor need to refer elsewhere, in order to appreciate fungal morphogenesis.

Contents

The first chapter is an overview of the evolutionary origins of fungi and the central role they played (and still play) in the evolution of life on Earth. The second chapter introduces hyphal growth, the essence of the fungal life style, and identifies features which are crucial aspects of morphogenesis. Chapter 3 summarises fungal primary and secondary metabolism, necessary here because adaptation of primary metabolism and exploitation of secondary metabolism are both critical to fungal morphogenesis. In chapter 4 the impact of physiology on morphogenesis is discussed, with the genetic components of differentiation and morphogenetic change being dealt with in chapter 5. The development of form and structure is the main theme of a lengthy chapter 6, and the ideas developed here are brought together and summarised in the final chapter 7.

Fungal Morphogenesis was well received on publication, indeed it appeared in the Top-ten Best Sellers list in the Professional Titles, Biology category produced by Yankee Book Peddlers for the 4th quarter of 1998. 

THE BOOK IS NOW AVAILABLE IN PAPERBACK AND YOU CAN ORDER AT A 20% DISCOUNT

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