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Debunking the New Morphology Author(s): E. J. H. Corner Source: New Phytologist, Vol. 65, No. 3 (Jul., 1966), pp. 398-404 Published by: Wiley on behalf of the New Phytologist Trust Stable URL: http://www.jstor.org/stable/2430058 . Accessed: 14/06/2014 00:08 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . Wiley and New Phytologist Trust are collaborating with JSTOR to digitize, preserve and extend access to New Phytologist. http://www.jstor.org This content downloaded from 194.29.185.109 on Sat, 14 Jun 2014 00:08:05 AM All use subject to JSTOR Terms and Conditions

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Page 1: Debunking the New Morphology

Debunking the New MorphologyAuthor(s): E. J. H. CornerSource: New Phytologist, Vol. 65, No. 3 (Jul., 1966), pp. 398-404Published by: Wiley on behalf of the New Phytologist TrustStable URL: http://www.jstor.org/stable/2430058 .

Accessed: 14/06/2014 00:08

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

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Page 2: Debunking the New Morphology

DEBUNKING THE NEW MORPHOLOGY

BY E. J. H. CORNER

Botany School, University of Cambridge

(Received 3 January I966)

A recent volume of Advancing Frontiers of Plant Sciences gives us the latest trend in this creed (Meeuse, I965). Professor Meeuse dissects the flower into bits and pieces of an ancestral branching system, recombines them, and pulls Magnolia out of the hat. This is his theory of Angiosperms-Past and Present. It reminds me of Melville's (I962) which I have already criticized (Corner, I963). I find this new version as misleading.

Neomorphologists turn their backs on the classical morphology. It is very hard to discover their reasons. Meeuse and Melville dismiss it summarily and imply that it is profitless. Yet one of its main products has been the explanation of the flower as a repro- ductive bud composed simply of stem and leaves. It is a most elegant theory, comparable with celestial mechanics because it assumes the minimum and derives the most. I am a classical morphologist proudly. In all my researches into flowers, fruits, and seeds, as well as stems and leaves, I have found more and more evidence for the classical theory. Once it was the brilliant inspiration of Goethe; now it is backed by an immense amount of structural botany. It is not kicked over by the New Morphology and it needs no championship, but this new book (Meeuse, I965) contains so many fallacies that in the modern rush for new outlooks they may appear real.

The disagreement between the new and the old morphology is one of approach. The old seeks to discover the unknown from the known and to infer the manner of evolution from its existing products. It is comparable with that part of geology which infers the past history of the Earth from its present rocks. The new morphology invents concepts, such as phyllome and telome, gonophyll and gynoclad, discovers them in fossils with all their imperfections, and reads them into modern plants as though the rocks had an infallible history of plant life. Its very approach seems to me unscientific.

For instance, as the leaf is such an important item of classical morphology, I set myself to understand as much as I could about the diversity in shape, structure, composition and development of the leaf in living plants. I have published little because my findings agree mostly with what has been published; this does not mean that they are appreciated. From chance passages I conclude that Meeuse, like Melville, has not permitted himself the same preparation when he dismisses the foliar nature of stamen and carpel. Meeuse quotes sympathetically from Bremekamp that 'the term "leaf" has outside the Phanero- gams no morphological meaning' (p. 36) and approves the conclusion that the idea of a sporophyll is equally meaningless (p. 41). But I have a clear idea of the meaning of the term 'leaf' for the dorsiventral appendage produced in phyllotaxis at the stem-apex, and I can recognize the leaf in mosses, ferns and seed-plants, even when it is merely microscopic or cylindrical: and I could never have learnt this from fossils. Furthermore I can recognize fertile leaves and distinguish microsporophylls from megasporophylls shamelessly. They are not postulates or concepts (p. 41), but things which I can cut off and show to others. I can weigh them and measure their growth, and experimentalists can

398

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Page 3: Debunking the New Morphology

Debunking the new morphology 399 displace them. When you come to think of it, it is remarkable how little palaeobotany could know of leaves on its own account. Phyllotaxis, however much inexplicable, is an all-pervading method of shoot-construction and its products, stem and leaf, are as truly homologous throughout the plant kingdom as any plant structures can be; we do not deny the homology of cells because some are haploid and others diploid. Meeuse, like Melville, seems oblivious of the importance of phyllotaxis as the means of making the angiosperm shoot, and the failure to relate their theories with it reflects their inadequacy.

Meeuse writes of the dichotomy of leaf-venation in Nelumbo as though it indicated great primitiveness and antiquity for the genus. Now equally good, if not better, dicho- tomous venation occurs in the very advanced and far from antiquated, epiphytic fig-bush Ficus deltoidea, of which more collections have been made in tropical Asia than of any other wild fig. Here the dichotomy is a sign of advancement in the sense that this species of Ficus has lost the normal control of making a fig-leaf. When apical growth disappears in leaves, midrib and side-veins relapse into dichotomy. Thus obtuse and retuse leaves generally show dichotomy of the midrib towards the distal end in the same way as many fern-leaves. The broad blade of Nelumbo, like the small blade of Ficus deltoidea, which also has a peltate variety, is advanced and has lost its power of extensive apical growth which rectifies dichotomy into a straight line. Through such apparently small mis- directions, arising from ignorance of classical morphology, most erroneous standpoints can be achieved. Correct them, and the new morphology wavers.

Meeuse, like Melville, deduces the primitive flower from fossil and gymnospermous reproductive structures: for Meeuse they are Bennettitalean and Gnetalean. But, with others who uphold the classical theory, I endeavour to deduce the primitive flower from the construction of living plants. Meeuse condemns this as preconceived. I regard it as scientific and based on facts, and consider his procedure to be preconceived and fictitious for there are no fossils or living gymnosperms which link up with angiosperms; hence there is no proof that any of these Bennettitalean or Gnetalean states have any ancestral connection with the angiosperm. It is he who forces the facts to fit a hypothesis. The classical, and scientific, theory infers from observation. Flowers, just as the structures of any group of organisms, show stages in progress, and these must be interpreted as evolu- tionary. The outstanding features of the flower are the reduction of its direct and primary apical growth as a bud and the substitution of this by indirect, secondary, intercalary, and basipetal growth; the result is the tubular construction with inferior ovary. It is this consequence, paralleled in many alliances of flowering plants, which enables the classical theory to reconstruct; and the reconstruction requires understanding of floral develop- ment. Here lies another big difference. Neither Meeuse nor Melville pay attention to the manner of development of flowers, as if one could understand anything living without knowledge of how it is made. It could be chalked up in large letters against their theories that the flower never develops in the manner in which they suppose it to have evolved. It should be chalked up in the largest letters in favour of the classical theory that the flower develops in the manner which the theory necessitates. Stamens and carpels develop as leaf-primordia in phyllotaxis and the transition from leaf to bract, sepal and petal continues in varying phyllotaxis to stamen and carpel. There is never any sign of a double origin for the carpel primordium as the theories of Meeuse and Melville demand; nor is there any sign of a double or multiple origin for the stamen primordium, though fasciculate and centrifugal stamens may give such a spurious appearance. This admission, by itself, condemns their theories. They remind me of the endeavour to find the homun- culus in the gamete because the preconception must be there.

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Page 4: Debunking the New Morphology

400 E. J. H. CORNER

In seeking to establish what the angiosperm is, Meeuse also omits the fruit. I maintain the obvious standpoint that the fruit, which forms and sends forth the seed, is one of the prime characteristics of the angiosperm to which the flower is a precursor. Thus Meeuse, like Melville, omits all the facts, deductions and inferences of the durian theory, and the word 'fruit' has merely trifling reference in his index. Instead, he picks out the aril, which is a fruiting feature of the seed, and ascribes it to a fossil ovuliferous involucre. This is not scientific logic, step by step, but jumping to a conclusion; it is the preconception which he strangely misapplies to the whole framework of the classical theory, as if botany for a century had been engrossed with fabrications of the mind. The aril must be evaluated from the post-fertilization development of the ovule; it is a problem of the fruit, not of its developmental precursor, the flower. And what do we find? In those very groups of flowering plants which Meeuse holds up as primitive and, therefore, nearer to his arillate-involucrate cycadopsid ancestor, namely Pandanaceae. Palmae, Araceae, Cyperaceae, Juglandaceae, Myricaceae, Urticaceae and Piperaceae, as well as Gnetales, there is no aril. On the classical theory, these families are far advanced with highly reduced flowers and indehiscent fruits; they are understandably exarillate. Here is the straw which shows the wind hard against this preconceptual theory of the anthocorm with androclads and gynoclads.

Meeuse is distressed with the apparently heterogeneous assemblage of plants under the name angiosperm. 'One only has to compare the humble daisy with an oak, a water- lily with a palm . .. to distrust the hypothesis of monophyletic descent' (p. 4). 'One need only compare a Ficus or a Betula with a Magnolia, Nymphaea, Polygonum, Areca, Poa, Carex, Arum and Orchis to agree that they certainly represent widely divergent mor- phological "types" ' (p. 2I). I find these statements remarkably naive. Classical mono- phyletic theory readily knits Urtica dioica with the giant stinging tree Laportea, the tiny Fatoua with a vast banyan or a geocarpic fig with naked ovary, Lemna with Amorpho- phallus, Cassytha with Lauraceae, Nardus with Ochlandra, or Lathyrus nissolia with Parkia, Dialium, and even Inocarpus. Yes, try to classify this great and common, um- brageous chestnut of the Western Pacific, Inocarpus, known to millions of villagers, which neither in leaf, flower, fruit nor seed is leguminous, and perceive the aberrations of Na- ture and the power of classical morphology! I fail to see why its ability to knit Nymphaea and Betula, or Ficus and Orchis, should be discredited. The differences are not more striking than between a whale and a marmoset or a butterfly and a scale insect. These comparisons aggrandize rather than minimize the classical theory.

Likewise Meeuse seeks to separate monocotyledons from dicotyledons phyletically, but some of his criteria are false. It is absurdly wrong to deny axillary buds in general to the vegetative parts of monocotyledons (p. 3I). I recall bamboos, branching pandans and dracaenas, suckering palms and bananas, and a host of branching rhizomes, bulbs and corms. It is wrong, also, to refer to the monocotyledons as branching dichotomously or trichotomously; they are distichous or tristichous with axillary branching, and the one exception appears to be the truly dichotomous trunks of Hyphaene. Thus superficialities creep in to obscure the rigid and clear thought of the classical theory.

On p. 47 of his work, Meeuse summarizes the evolution of the angiosperms as progress in anatomy, pollen structure, gametophytic reduction, double fertilization, and com- pleted seed-embryo. He seems unaware of the versatility of leaf, stem and cambium by which they have built the majestic forests of the world. On p. I4, 'Corner was certainly right in drawing our attention to evolution in the tropics', but with what avail? The versatility of the leaf has produced the pitcher of Nepenthes, the frond of the palm, the

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Page 5: Debunking the New Morphology

Debunking the new morphology 40I

tendril of Lathyrus, and the thalloid Podostemonaceae. The modification of the sporo- phylls into stamens and carpels, then staminal tube and syncarpous ovary, are small events compared with the vegetative transformations. To swallow these and to cavil at the classical theory of the flower is to strain at a gnat.

What puzzles Meeuse, as they puzzled Eichler, Engler, Wettstein, de Vriese, Lam and many other European botanists, are the minute and, often, anemophilous flowers. It seems incredible that they are simplifications of the classical Magnolian prototype, yet the study of the whole extent of such enormous alliances as Leguminosae, Sterculiaceae, Euphorbiaceae, Sapindaceae, Moraceae, Gramineae, Scitamineae and even Palmae show step by step in living plants the gradation from relatively large, classically constructed flowers to minute unistaminate and often unisexual and apetalous flowers aggregated in anthocorms or catkins. When the flowers of any one alliance are studied developmentally it can be seen how these minute flowers are principally neotenic bud-products. So there are steps in neoteny from Delonix to Ceratonia, from Ravenala to a cleistogamous Marantaceous flower with half an anther, from Jatropha to Macaranga, Sapium and Euphorbia, or from Morus to Artocarpus and Ficus. I recommend the study of these vast families or orders to those who refuse to acknowledge the general trend from the Magnolian type to the Piper-type. It puzzles me why exponents of the evolution of the flower close their eyes, or their minds, to this enormous body of living evidence in preference for a few imperfect fossils; the neglect renders their theories resoundingly hollow.

I have frequently wondered to what speculation the discovery of well-fossilized figs of the geocarpic Ficus uncinata, so common in south-east Asia, might give rise. Here are bracteate cupules, 2 in. wide, borne on bracteate twigs, and filled with cupulated ovules for the female flowers have no perianth; those with cupulated stamens (one to a cupule) would show no embryos, being gall-figs (which the fossil would not disclose), while those without male flowers would contain embryos. They would appear as magnificent cupu- lated anthocormic supervasculated Caytonialean protocarps containing arillate ovules, as shown in Figs. 2 and 4 of Meeuse's volume. But, when the whole living gamut of Ficus and Moraceae is studied in the detail which is never given to palaeobotany, the derivation of F. uncinata from the racemose, extrovert inflorescence and reduced Magnolian type of flower of Morus is apparent; and, strange to say for the anthocorm theorists, this conclusion is universally admitted. Classical theorists expect that in a similar way it will be shown how the Piper flower, the Engelhardtia flower and the Cyperaceous flower have been derived; they expect, even, that with advance in experimental ability the less derived states of these reduced flowers will be recovered in the experimental laboratory. While we cannot now positively disprove the fictitious sketches relating Caytonia and Gnetum with amentiferous flowers, the disproof will come through genetical biochemistry. It is noteworthy that teratological flowers revert to effects of the Magnolian type and not to anthocorms; and what are teratological effects but the outcome of that transference of function which has been the main process of floral evolution?

The odd fact is that no textbook explains what the syncarpous ovary really is. Hence it is easy to delude oneself with sketches of Cyperaceous or Pandanaceous flowers (Figs. 9 and I3 of Meeuse). There is such a vast variety of flowers that many sorts can be picked out eclectically to fit as many theories. The scientific method is to proceed by comparative morphology, including ontogeny, in natural alliances and to become objectively informed of the process of development and structural change. It is what the palaeobotanists would do if they could. Thus the classical theory of the flower, which

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Page 6: Debunking the New Morphology

402 E. J. H. CORNER

began certainly with poetic and philosophic intuition inspired by the regular beauty of the flower, has built around it the firm edifice which palaeobotany is without; indeed, I wonder if it would be possible to interpret any plant fossil without a knowledge of living plants. Nevertheless, Meeuse and Melville denounce the classical morphology as tra- ditional dogma; in its place are statements which begin 'I believe. . . 'or 'In my opinion ... ', so detrimental to the name of botanical science. They support these beliefs with outline sketches of superficial structures, tentative or derived from the suggestions of others, when we should be thinking of the structures as the outcome of cell-division. To me this is doodling. Thus (p. 7I and Fig. 6) Meeuse believes that the ovary of the common weed Leucosyke (Urticaceae) is really comparable with a very primitive cupulate (chlamydote, to make it more mysterious) ovule of gymnospermous type, and would have us believe that this super-abundant plant, which we hack out of our way with knives, is a missing treasure. What is entirely omitted in the academic aspect of this plant is the comparative morphology of the ovary in Urticales which shows that the unilocular ovary of Leucosyke is derived by abortion from the bilocular ovary, as in Morus, and that the lateral anatropous ovule of the Moraceae becomes displaced to the basal orthotropous ovule in Urticaceae. It is erroneous, if not actually misleading, to extract and present in isolation the intricate and simplified structures of such modern and advanced plants as Leucosyke. We might as well present the minute unistaminate flower of Artocarpus as the primitive cupulate androclad.

The pre-flower of Meeuse's imagination had a slender elongate bracteate axis bearing slender ebracteate but axillary branches-axillary through design rather than as a concession to classical morphology-on which were set with internodal spacing either stalked ovules or sessile anthers (male synangia, to be precise). The whole is the antho- corm with gynoclads or androclads (Fig. z). In Fig. 3, the anthers become stalked. There is no explanation how a branch-system has internodes without nodes, that is without leaves or bracts, but I suppose such details of construction can be ignored by neomorphologists. The male organs (even genitalia) in Fig. 3 resemble normal stamens, but the author does not realize that the ordinary stamen is a very derived structure in which the filament is an intercalation homologous with an intercalary petiole in a basi- petally developed leaf. It would appear from the illustrations that these stalked genitalia are homologous with the bracts of the anthocorm and, therefore, even sporophylls, but I doubt if this is Meeuse's intention. I must conclude that this quaint structure, called the anthocorm, could not have been developed ontogenetically in the manner in which the parts of living flowering plants develop, and it is useless, therefore, for the classical morphologist to pursue the matter further; if the upshot of some extinct race of seed- plants, it could not have lead to modern flowering plants. Yet, I will give two more reasons for rejecting the theory.

This anthocorm is supposed to have contracted, not unexpectedly, by some sort of loss of 'internodal extension' into its bud as a capitulum. This, by loss of bracts between the anthoclads and by reduction of the anthoclads to one anther gave the normal androecium below the gynoecium. In the female part, however, besides the contraction of the gyno- clads, there was a synthesis or fusion between them and the attendant bracts-hence, no doubt, their convenient axillary status-and the conjoint structures made the closed carpel in much the same way as Melville's gonophyll behaved. Thus, with still elongate axis, but contracted clads (branchlets), the Magnolian flower is derived in Fig. I5. (If I turned a fig inside out and removed a few bracts, I could imagine a Magnolian flower also.) Admittedly Meeuse describes this as a tentative explanation of Magnolia, but even

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Page 7: Debunking the New Morphology

Debunking the new morphology 403 if this is true for all his explanations it is difficult to see for what purpose they are tenta- tive: they mystify rather than help, and in the absence of any compelling physiological trend such series can be read in any direction. The analogy among living plants is the capitulum of Euphorbia. Here, however, terminal flowers contract in a cymose panicle to give many unistaminate male flowers, like Meeuse's primitive stamens, around and below a single terminal female flower. I am not sure that there is any example among living flowering plants of a racemose structure with lateral flowers condensing to give a terminal gynoecium. There is an important physiological or morphogenetic difference between the axillary and the terminal positions, and botanists of the classical school have a right to ask why Meeuse reverses the natural trend and makes lateral structures become terminal.

Then, as with the gonophyll, the question is unanswered how the gynoclad or raceme of ovules fuses with the conveniently attendant bract; clads should not fuse with phylls in these neomorphological constructions. When I read such facile statements written without explanation, I realize at once that the author does not understand how the so- called fusions of parts-really, substitutions-arise by intercalary growth accompanied by transference of function to the insertion. It is a favourite dodge of flower-theorists to fuse parts, as if by squashing the bud; it is so fallacious that one is shocked. If the real process of 'fusion' were understood, the strength of the. classical theory would be revealed. Starting with the classical and demonstrable apocarpous flower with free parts, all other kinds can be derived in the manner in which the flower grows. Then, as men- tioned, teratological variations are understandable as abnormal transferences of function. I repeat that these abnormalities always support the classical theory, never the hypo- theses of neomorphologists, for the simple reason that flowers never work by the un- natural steps that they devise. Teratology will become a fascinating subject, as genetical biochemistry and morphogenetics progress, but it is at present still rather too shockingly classical.

To prop the theory, Meeuse introduces fasciculate stamens and centrifugal stamens. They could be, and they should be, contracted androclads. Unfortunately, fascicles of stamens always develop as separate anther primordia in normal phyllotaxis, but at some subsequent stage they are raised on mounds of basipetal intercalary growth into fascicles. There is never an outgrowing axis from the floral receptacle along which the anthers develop, as the anthoclad theory would demand. Hence Meeuse distorts the facts which are easily explicable on the classical theory. He is obsessed, too, by the idea that centrifugal stamens, themselves often fascicled, must represent innate branches because the vascular supply to them branches inside the stem from a common initial; they have sunk into the floral axis, but how? This was Wilson's notion when he studied the sta- mens without enquiring how they developed. However, there are no more androclads in the centrifugal androecium than in the centripetal. In the centrifugal there is another instance of the universal floral process of intercalary receptacular growth; it happens in this case to enlarge the interval between petals and stamens during the phase of anther- making and causes new anther-primordia to develop in reversed phyllotaxis below, or external to, the first stamens. It is certainly peculiar, but that does not mean that it condemns any or all and every theory of floral evolution. It has to be accepted in much the same way as entire or simple leaves may continue to grow intramarginally by inter- calary growth and add extra loops of dichotomous veins, e.g. lettuce leaves. I wonder that Meeuse has not been struck by the resemblance between centrifugal stamens and the cen- trifugal unistaminate male flowers of Gnetum.

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Page 8: Debunking the New Morphology

404 E. J. H. CORNER

I pass over many absurdly dogmatic and highly questionable statements, interspersed authoritatively through the text, such as 'primarily unisexual flowers' in palms and Monochlamydeae, 'primitive groups' for Cyperaceae, Pandanaceae, Dilleniaceae, Laura- ceae and Araceae, the separate origin of monocotyledons from dicotyledons, and the 'concept of the flower' as if it were not real. And I come to my last point which tells the total inadequacy of the anthocorm theory.

In all the sketches illustrating his theory, except that of Magnolia, which we know so well, Meeuse shows the axis of the anthocorm as a line. If permissible schematically, it is unrealistic and obscures the important relation between primary stem-size and the nature of the appendages; this is what I have represented as pachycauly with megaphylly against leptocauly and microphylly. It is far from clear that a primitive pachycaul would produce anthocorms which are essentially leptocaul. Meeuse thus represents his primi- tive flowering plants as leptocaul with small simple leaves. From them, I suppose, he would derive the pachycaul, but he makes little reference to the vegetative nature of the primitive flowering-plant. The durian theory dealt explicitly with this problem because it has prime importance in understanding how the flowering plant evolved as forest. If there is any truth in Meeuse's theory, which I doubt, then he will have to ex- plain how leptocauls evolved into pachycauls and where there is evidence for such a conclusion. All the flowering plant families that I have examined point to the reverse conclusion. From fruit, flower, carpel, stamen, seed, leaf, stem and indeed root, the evidence is that pachycauly with overall massive construction preceded leptocauly and slender construction. The problem of the flower cannot be solved in abstraction from the rest of the plant, and this is where the modern theories fail which so sanctimoniously extol themselves as the new gospel. 'Only ideas emerging from the New Morphology, i.e. from a dynamic approach of morphological problems, could pave the way to a solu- tion of the problem of Angiosperm evolution so that one can now . . . discard all alterna- tive hypotheses and speculations "to get down to some good solid work" ' (p. I75).

What is new in this Morphology is ignorance of the old, and such conceit is the reced- ing frontier of plant science.

REFERENCES

CORNER, E. J. H. (I963). A criticism of the gonophyll theory of the flower. Phytomorphology, 13, 290. MEEUSE, A. D. J. (I965). Angioxsperms-Past and Present. Advancing Frontiers of Plant Sciences ii.

Institute for the Advancement of Science and Culture, New Delhi. MELVILLE, R. (I962). A new theory of the angiosperm flower. I. The Gynoecium. Kew Bull., I6, i; II.

The Androecium. Kew Bull., 17, I.

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