Makers of British botany/William Griffith 1810—1845

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Early training—medical appointment under the East India Company—his travels—the magnitude of his collections—his method of work—results of researches mainly published posthumously—the ovule and fertilisation—Santalum—Loranthaceae—Balanophora—Avicennia—his gymnosperm work illustrated by Cycas—discovery of the pollen-chamber—Rhizocarps and Liverworts—pre-Hofmeisterian work—Griffith's relation to his times.

It might have been assumed that all the names of British botanists whose work has been or is to be considered in this course of lectures would have been familiar to their successors of to-day, even if their works were too often neglected for the last words of scientific progress in a summary of literature. The question has however been put to me by more than one botanist in the last month or two, "But who was Griffith?" That this should be possible seems in itself ample justification for including his name in this list of British botanists.

For Griffith has claims to be regarded as a great botanist. It is true that he failed to break through the limitations of his time and period—that he left no new and more correct general views to modify the science. But this is true of all his contemporaries, indeed it is true of most botanists. To recreate the department of a science in which a man labours requires a combination of ability and fortunate chance that is given to few.

Griffith had the ability, the power of independent observation, the readiness to speculate, the careless prodigality of labour. He did not however, in the fraction of an ordinary working life that

Plate XV

fate allowed him, attain that insight into more correct comparison of the plants whose morphology he studied which would have acted quickly on the mass of first hand observation he possessed.

It is well to be clear at the outset that it is the personality of William Griffith, his important detailed contributions to botany, and his achievement as a great working morphologist of his time that will interest us to-day—rather than his general views or any influence of these on the progress of botany. Griffith had the advantage or disadvantage of botany being his private study and not his profession. The motive force of his career was however his love of scientific work for its own sake.

William Griffith was a London botanist. He was the son of a London merchant, born on March 4, 1810, at Ham Common. Having finished school he began to prepare for the medical profession and was apprenticed to a surgeon in the West end of London. About 1829 he commenced attendance at the classes in the newly established University College. He had earlier in life shown an interest in natural history but was now specially devoted to botany. He attended Lindley's lectures, and also studied medical botany under Mr Anderson at the Apothecaries' Garden in Chelsea. There he obtained the Linnean Gold Medal given by the Society of Apothecaries. At this time also he was a frequent visitor to Kew Gardens where he was on good terms with the head gardener and also came under the influence of Mr Bauer the great botanical draughtsman of his day. Griffith was never tired of expressing his admiration for Bauer as an accurate observer. During his vacations Griffith made botanical excursions in England, carrying his light baggage and his equipment for collecting plants.

That the training that Griffith received in botany in the London University of that date was a sound one is shown by his power of facing the most various problems when cast on his own resources immediately at the close of his University training. The soundness of his training is further shown by the small pieces of original work he had published before leaving England at the age of 22. Not only had he made some of the illustrations for Lindley's Introduction to Botany and had described the flower and the structure of the wood of Phytocrene gigantea in Wallich's Plantae Asiaticae Rariores, but (a noteworthy indication of his interest in Cryptogams at this time) he had supplied an account of the structure and development of Targionia hypophylla to be appended to Mirbel's classic monograph on the anatomy and physiology of Marchantia polymorpha—published in 1832.

His medical studies finished, Griffith sailed from England in May 1832, he arrived at Madras in September and was appointed Assistant-Surgeon on the Madras establishment in the service of the East India Company. His scientific work was done in the intervals of a busy life. Only a man of great energy and enthusiasm and possessed of great powers of physical endurance could have done the work that Griffith crowded into the years, between his landing in India and his death at Malacca before the age of 35 on February 9, 1845. This time was all spent in the East Indies—he never returned to England.

Deferring for the moment consideration of his scientific work we may take a general survey of Griffith's movements during his working life and of his labours as an explorer and collector.

After spending some months in the neighbourhood of Madras, he was situated for more than two years at Mergui and collected extensively in Tenasserim. He was recalled to Calcutta in 1835 and attached to the Bengal Presidency in order to be sent with Dr Wallich and Mr M'Clelland to visit and inspect the localities in which tea grew wild in Assam. Griffith's full report on this enquiry led to the important economic conclusion (based largely on a critical comparison of the Assam flora with the flora of tea-growing regions of China) that tea might be successfully grown under the conditions in Assam and similar districts of India. When the other members of the expedition returned Griffith was detained in Assam, where he remained during the whole of 1836, making a successful expedition into the Mishmee mountains only once before visited by a European.

Early in 1837 Griffith, accompanied by only one servant, set off on an exploring expedition through the very disturbed country of Burmah towards Rangoon. All news of him ceased, or rather his assassination was credited by the Government and reported in the newspapers, when in June he re-appeared, ragged and travel stained, in Calcutta. He had explored down the Hookhoom (Hokong) Valley and on to Ava, and had then proceeded more rapidly by river to Rangoon, conveying his collections with danger and difficulty.

Appointed Surgeon to the embassy about to start for Bhutan, he filled up the intervening two months by again going to the Khasi hills to collect. He then accompanied the expedition to Bhutan, traversing over four hundred miles of the country and returning to Calcutta in June 1838. Here he spent the next few months arranging his collections and also studying the plants of the suburbs.

In November he joined the army of the Indus and accompanied it in its whole march. He remained another year in Afghanistan making various expeditions in the country and into the Hindoo Koosh. He returned, after visiting Simla and the Nerbudda, to Calcutta in the middle of 1841.

Griffith then proceeded to Malacca where he had been appointed Civil Assistant-Surgeon. He remained only a year, but long enough to appreciate the great interest of the district for his botanical work and to complete some important observations. He collected the plants of the province and also visited Mount Ophir.

Recalled to Calcutta, he took charge of the Botanic Gardens and also lectured to the medical students during Wallich's absence from August 1842 to August 1844, pressing forward reforms in the gardens and using his opportunity for scientific observation. On Wallich's return Griffith remained for some months longer in Calcutta continuing his work, married in September, and returned to Malacca in December full of hopeful plans for scientific work there. He had barely arrived at Malacca and begun work than he was seized with a fatal illness and died on February 9, 1845.

It has been necessary to consider in some detail the rapid movements of Griffith's life in the East in order to fully appreciate the difficulties under which his large amount of scientific work was accomplished. The twelve years of his official life were filled with professional duties, difficult and dangerous exploration, management of the Botanic Gardens, and the labours entailed in making and caring for extensive collections. It would not have been surprising had Griffith, in spite of his attainments, contributed nothing to scientific botany beyond rendering these collections available for other workers. He estimated his collection of plants at more than twelve thousand species; and on his travels he did not neglect other collections of interest. Insects obtained by him are described, he collected the birds and fish in every district he visited; indeed he was a keen fisherman and must have thrown a fly in many a stream that had not been fished before, combining sport and science.

Griffith's collections were made with the definite purpose of enabling him, when he had leisure, to produce a general account of the Indian flora on a geographical basis. His methods of collecting were most enlightened and subserved his work as a morphologist and a student of the conditions of occurrence of the plants, not merely of formal systematic botany. The journals he kept on all expeditions are full of references to the occurrence of the plants met with. He often adopted a plan of roughly mapping each day's route and indicating the plants and associations of plants, along the line of march. I wonder if modern ecologists know of these records made long before ecology was invented?

Whenever possible he seems to have examined the morphology of the living plants, and he fully realised the value of preserving portions of the plants in spirit for future examination instead of relying on herbarium material.

This quotation from a letter to Wight (then Superintending Surgeon of the Madras Service), with whom Griffith kept up a most interesting and friendly correspondence, from which I should like to quote largely, may give an idea of his point of view and also show how he looked forward to returning to Malacca:—

"If ever you go to the place of Podostemon endeavour to get some germinating or at least very young plants. I can fancy how an Acotyledonous plant gets a stem but how a Dicotyledonous plant loses it, and becomes as some of them do, mere discs spread over rocks is another thing. Then again where are their roots? How opposed to late ideas of the absolute distinction of the three great divisions. Also please to take a bottle of spirits, and deposit specimens in it. I shall not be very sorry to get back to Malacca, this is a delightful place truly, but one is interrupted, and the lectures at the Medical College consume much time. For botany no place can exceed Malacca."

And again,

"What a business it will be to settle the types of the families from which the names must eventually be taken; this will never be done by dried-plant botanists; but by examination of development, which I am convinced will alone give the key."

As to Griffith's methods of work, we learn from a memorial notice of him by Mr M'Clelland that whenever possible after the business of the morning was finished the rest of the day was devoted "to the examination and dissection of plants under the microscope, drawing and describing all peculiarities presented." "Even on his death-bed his microscope stood beside him with the unfinished drawings and papers and dissections of plants on which he was engaged the day on which the fatal symptoms of his disorder came on."

All his work shows the same characters of direct individual observation and interpretation of the facts before him, repeated examination of the same point, and almost a prodigality of labour in recording his observations in drawings. At first under the influence of Robert Brown, he used the simple microscope with triplet lenses, but later he employed the compound microscope and in the year before his death writes hopefully of ordering a first-rate microscope when he obtains the arrears due to him from the Directors.

Griffith's high attainments were appreciated by the distinguished circle of English botanists of his time with whom he corresponded. Mr Solby, to whom he always sent home his papers for submission to the Linnean Society; Robert Brown, to whose work he constantly recurs with admiration, and whose judgment he trusted absolutely; Lindley; Sir William Hooker, who looked forward to his being settled permanently in charge of the Calcutta gardens, and Dr Wight may be named.

I may quote from a letter addressed to Griffith by von Martius of Munich, since it couples his own opinion and that of Robert Brown. "He (Brown) agrees with me in appreciating your spirited and enlightened investigations, and I now more than ever look forward to you as his successor—as the standard English botanist."

Only an outline of the nature of Griffith's scientific work with some details on selected subjects can be attempted here. His published works in the Transactions of the Linnean Society and elsewhere, important as they are, represent only a small fraction of his observations. But the wisdom and liberality of the East India Company has put us in possession of his unpublished notes and drawings (bequeathed with his collections to the Company) in the posthumously published volumes of Notulae ad plantas Asiaticas with the accompanying sets of plates. Though his papers were not ready or intended for publication in this form and suffer from having had to be arranged by another hand, they afford, together with his published work, a particularly good picture of how the problems of morphology and classification presented themselves to a keen investigator at this time.

Of his purely systematic work I shall not speak at length. In addition to smaller papers the most important contribution was his illustrated monograph on the Palms of British East India. In the Notulae numerous species are described and figured nearly always with reference to the morphology and physiology of the parts concerned. It is his investigations made with direct reference to morphology and reproduction that claim our attention most. In dealing with them it is convenient to treat of the main questions to which he directed his attention rather than of the separate papers. I shall call attention first to his work on the flower and on fertilisation in a number of plants, then to his observations on Cycas, and lastly to his work on the Cryptogams.

Interest in the structure of the ovule and the nature of fertilisation was widespread at the time Griffith worked. A few years previously Robert Brown had laid the foundations of the scientific study of the ovule and the behaviour of the pollen tube, and during Griffith's time the papers of Schleiden, which extended the comparative study of the ovule and advanced the important though erroneous view that the embryo originated inside the embryo-sac from the tip of the entering pollen tube, were appearing. Schleiden's text-book did not appear until too late to be known to Griffith. His interest was keen on continuing the work, that Brown had begun, on plants that only a resident in the tropics had the opportunity of studying properly, and the first volume of the Notulae, with the accompanying Icones, and the more systematic volume on the Monocotyledons and Dicotyledons contain his unpublished observations on the ovule and flowers of many plants.

His first paper in the Linnean Transactions was on the ovule of Santalum. Griffith observed and rightly interpreted the free prolongation of the embryo-sac from the nucellus, and described the application of the pollen tube to the summit of the embryo-sac, the development of the endosperm, and the origin and development of the embryo. He also recognised and figured the great prolongation backwards of the embryo-sac as an empty, absorbent caecum. At first he left the origin of the embryo doubtful, while recognising the advantages of the exposed embryo-sac for settling the question, but later he decided in favour of Schleiden's erroneous view that the embryo developed from the tip of the pollen tube. Griffith also examined the ovules of Osyris recognising the corresponding facts.

Comparison with the figures of Santalaceous ovules in Guignard's later work will serve to show both the magnificent accuracy in observation of Griffith and the limitation, running through all the work of the time, of not recognising the contents of the embryo-sac before fertilisation.

The Loranthaceae was another family on which the development of the embryo-sac and the processes of fertilisation and development of the fruit interested Griffith specially. Not only did he send his results home to the Linnean Society in two papers, but his descriptions and figures of all the species described in the Notulae take account of these morphological and developmental facts. He traced the development of the cavity of the ovary and regarded the ovules as reduced to their simplest expression—to an "amnios" or embryo-sac. And he observed the extension of the embryo-sacs up the style and the union of the pollen tube with the tip of the embryo-sac. His further description of the development of the embryo, endosperm and fruit is wonderfully exact if we allow for his regarding the long suspensor bearing the embryo as derived from the pollen tube growing down through the long embryo-sac.

Griffith thus recognised all the main peculiarities of Viscum and of Loranthus subsequently described more in detail in European species by Hofmeister (whose analysis of Griffith's work in 1859 is a great testimony to its accuracy) and later by Treub in the tropical species which had been studied by Griffith.

The Balanophoraceae was another group, on which Griffith made pioneer investigations. He collected and examined all the species he met with, partly from the systematic interest in supporting Robert Brown's objection to Lindley's class of Rhizantheae, but still more from his interest in the details of their reproduction. An examination of the plates from his memoirs, only published after his death, in the Linnean Transactions will show how fully he was aware of the structure of the archegonium-like female flower of Balanophora; of the relation of the pollen-grains and pollen tubes to it; and of the appearance of the endosperm which he mistook for the embryo. Throughout he compares the structure with the pistillum (archegonium) of Bryophyta.

Thus in the Balanophoraceae also Griffith laid the foundations on which the work of Hofmeister, and more recently that of Treub and Lotsy follow.

When at Malacca Griffith interested himself among many other problems in the ovule and the development of the seed of Avicennia. He had previously paid attention to the viviparous embryos of other Mangroves. This piece of work, when compared with Treub's re-examination of Avicennia, brings out so clearly Griffith's accuracy, so far as his means of observation allowed him to go, that we may look for a moment at how these two investigations, separated by forty years, compare.

Griffith recognised the development of the embryo-sac in the nucellus of the ovule which he took to be naked, missing the very slightly indicated integument. He followed the pollen tube to the tip of the embryo-sac and the development of the endosperm in its upper portion, where the embryo appeared. He saw the growth of the endosperm leading to its complete protrusion from the ovule and inverting the embryo so that its cotyledons point to the surface. Further he saw the long, empty, absorbent caecum grow out from the hinder end of the embryo-sac into the massive base of the young seed.

This account is substantially correct in all its facts, and Treub's work adds to it the cellular details of the origin of the embryo-sac, the setting apart of the endosperm cell to grow into the haustorium, and the details of segmentation of the embryo.

Such vivid, accurate, description of strange facts, when previous knowledge gave no clue, is in itself no mean scientific achievement.

To sum up Griffith's work on the morphology of the reproductive organs of the Angiosperms we see that he added many important facts and gave correct descriptions of what still remain among the most anomalous ovules and embryos. His methods did not enable him to distinguish clearly the contents of the embryo-sac, and he accepted and confirmed Schleiden's erroneous view of the origin of the embryo. But this hardly detracts from the directness and consequent value of all his observations.

Turning now to the Gymnosperms, we find again that Griffith devoted much attention to those forms that from his residence in the tropics he was in a position to study with most advantage. He describes in the Notulae his observations on the ovules and pollination of various Coniferae and Gnetaceae. But we may concentrate our interest on his work on Cycas. The rough structure of the young seed had already been described by Robert Brown who had recognised the gymnospermy of the group.

But Griffith's descriptions and figures are much more accurate—are indeed far in advance of those of much later observers—and add greatly to our knowledge of this plant. These two figures (pl. XVI) will speak for themselves and show how clearly Griffith had grasped the morphology of the Cycadean ovule, how faithfully he delineated the details, and how he sought in progressive development to throw light on the structure. He added to the previously imperfect description of the ovule an accurate account of the pollen chamber, and the proof that pollen grains entered and filled it. Further he followed the germination of the pollen grains, not merely recording the fact that the tubes penetrated the nucellus all around the pollen chamber, but ascertaining in how many days the tubes were put forth. His fullest description is unfortunately displaced in the Notulae under the heading of Thuja, but it is clear that it refers to the Cycas figured on the same plate as that plant.

From what has been said of the nature of Griffith's work on the ovules, both of Angiosperms and Gymnosperms, the complete omission of his name in recent works on the two groups that are in constant use is at least noteworthy.

Griffith was specially interested in the study of Cryptogamic plants. In a letter to Wight he says "I would like to be out with a work on Indian Cryptogamia of higher forms; so much so that if I see no chance of my succeeding to the Gardens, I intend sending away all my other collections, and devoting myself to this object and general development, which is obviously the keystone of the arch."

He left Algae and Fungi (with the exception of the Characeae) alone, and it is his work on the Bryophyta and Pteridophyta that concerns us. For information on his views on these plants we are dependent on his paper on Salvinia and Azolla and on the Notulae, put together as I have said from his notes after his death, and not intended for publication in this form. But there is no difficulty in getting a clear grasp of his point of view. This was a mistaken one—an attempt to bring into line the reproduction of the gametophyte of Bryophytes, the sporophyte of Vascular Cryptogams, and the flowering plant with its flower and fruit. It is easy to be wise after the event. In these comparisons Griffith belonged to his time with a much wider field of personal observation than most possessed.

We must bear in mind that at the time when Griffith worked no idea of the sexual and asexual alternating generations in

From Griffith's NotulaePlate XVI

Median section of the ovule of Cycas Nucellar apex of Cycas with pollen chamber
and pollen grains

Pteridophytes had been gained, although the prothallia had been observed preceding the growth of the plant in Equisetum and Ferns. It was not till some years after Griffith's death that fuller facts as to the sexual organs were obtained and led to the right comparisons.

Griffith's work on the Bryophyta shows the same power of observation as that on the ovule, but the difficulties due to imperfect instruments are more evident. His views on reproduction were here, however, clear, since the development of the capsule was definitely related to the fertilisation of the pistilla (archegonia) by the substance formed in the anthers. His figures indicate how much he saw, and how here also he sought in development the interpretation of mature structure.

His early interest in the Liverworts, especially the Marchantiaceae, continued, and all the forms he collected were carefully examined and figured with his usual accuracy.

One of the Liverworts Griffith described may be taken as an illustration to this part of our subject on account of the interest of its re-discovery and re-description in 1910 by Goebel. This is a plant collected in Assam and named Monosolenium tenerum. This Marchantiaceous plant is described as having no air-chamber layer, as bearing sessile, dorsal, antheridial receptacles, and terminal, shortly stalked archegoniophores with one ventral groove in the stalk. A single archegonium—later capsule—is found in each of the half-dozen involucres. Spores and irregular bodies were found in the capsule.

Recently Goebel had two tea-plants sent home from Canton. They died, but he kept the soil moist on the chance of germinating seeds. Among a number of other plants there turned up a new Liverwort. On examination this proved to be Griffith's Monosolenium—all types of which had been lost—a most interesting form related to the Corsiniaceae.

In the Mosses and the Liverworts generally Griffith was clear on the development of the capsule or fruit following on the impregnation of an archegonium. But in Anthoceros, while he recognised the antheridia he was not clear as to the sunken archegonia, and regarded the capsules as arising by impregnation of unrecognisable spots on the young frond or thallus. He observed however the indication of the canal of the archegonial neck above the young capsule.

Analogy with Anthoceros confirmed him in his views on the reproduction of ferns. Here he spent much labour in considering the view, originally due to Hedwig, that the ramenta were male organs by the effect of which the sporangia developed. Griffith saw that if this was so, since the sporangia are initiated very early, the only time to search for the male organs was in the very young stage of the leaf. On examining such young leaves he found the terminal cells of the young ramenta very prominent and formed the working hypothesis that they were the male organs. But he stated this cautiously and was well aware how imperfect his means of observation were.

The whole line of work brings vividly before us how cryptogamic the Cryptogams were at this period.

Without attempting to survey Griffith's views on the various groups of Vascular Cryptogams, a word must be said of those on Salvinia and Azolla, on which he published a long paper in addition to the other descriptions and figures in the Notulae. His observations bear on the development of the sorus and sporangium, but he dismissed the microsporangia as abortive or imperfectly developed structures. (I may note in passing that the study of their development led him to regard the microsporangia of Isoetes in the same way.) He dwelt on the similarity of the sporangium and indusium of Azolla to a gymnospermus ovule, and regarded the filaments of Anabena seen penetrating within the indusium as probably the fertilising bodies in this naked-seeded cryptogam.

Thus with a large amount of fresh and original observation Griffith was on wrong lines in his general views and comparison—he classed the higher Cryptogams in his Notulae as







Griffith's general views of the reproduction of all the Vascular Cryptogams was necessarily wrong, since the prime clue of the recognition of the prothallus and plant as distinct had not been found. In this connection his figuring young plants of Equisetum attached to prothalli is interesting. In some speculations concerning the embryology of Loranthus he came, by a wrong line of approach, within touch of the right comparison, when he compares the endosperm to the confervoid green growth (i.e. the prothallus) at the base of the young plant of Equisetum.

It is idle to speculate on what might have happened had such a wide observer as Griffith chanced on the clue. In this respect he was of his time as most are. The man who put the industrious but blind gropings of this period in morphological botany straight, both as regards the development of the embryo and the comparative ontogeny of archegoniate plants was Hofmeister, and like all exceptional men he belonged to the new period created by him.

The great advantage of this course of lectures seems to me to be that it approaches the study of the history of botany in the right way; for progress in our science has been the result of individuals rather than of schools. The consideration of the work of Griffith from 1832 to 1845 is a vivid illustration of the condition of morphological botany in the earlier portion of the period, surveyed in one of the chapters in Sach's History under the title of "Morphology and Systematic Botany under the influence of the History of Development and the knowledge of the Cryptogams." These two subjects were always before Griffith.

The interest of the personality of William Griffith and of the work he accomplished in his tragically short life is obvious. Not less so is the way in which that work was done inside the limitations of his period. We, who are still gleaners in the field that Griffith and his contemporaries cleared and Hofmeister marked out and tilled, are probably just as incapable of conceiving the future developments of morphology.