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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

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This historic 1885 book edited by Foster and Sedgwick is the first of Francis Balfour's collected works published in four editions. Francis (Frank) Maitland Balfour, known as F. M. Balfour, (November 10, 1851 - July 19, 1882) was a British biologist who co-authored embryology textbooks.

The Works of Francis Balfour

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.

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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

The Works of Francis Balfour

Francis Balfour (1851-1882)
Francis Balfour (1851-1882)

Volume I. Separate Memoirs

M.A., LL.D., F.R.S.,

Fellow Of Trinity College, And Professor Of Animal Morphology In The University Of Cambridge.

Edited By M. Foster, F.R.S.,

Professor Of Physiology In The University Of Cambridge ;


Adam Sedgwick, M.A.,

Fellow And Lecturer Of Trinity College, Cambridge.

Macmillan And Co.


The Right of Translation is reserved.


Upon the death of Francis Maitland Balfour, a desire very naturally arose among his friends and admirers to provide some memorial of him. And, at a public meeting held at Cambridge in October 1882, the ViceChancellor presiding, and many distinguished men of science being present, it was decided to establish a ' Balfour Fund ' the proceeds of which should be applied : firstly to maintain a studentship, the holder of which should devote himself to original research in Biology, especially in Animal Morphology, and secondly, ' by occasional grants of money, to further in other ways original research in the same subject '. The sum of ^8446 was subsequently raised; this was, under certain conditions, entrusted to and accepted by the University of Cambridge; and the first 'Balfour student' was appointed in October 1883.

The publication of Balfour's works in a collected form was not proposed as an object on which part of the fund should be expended, since his family had expressed their wish to take upon themselves the charge of arranging for a memorial edition of their brother's scientific writings.

That edition, with no more delay than circumstances have rendered necessary, is now laid before the public. It comprises four volumes.

The first volume contains, in chronological order, all Balfour's scattered original papers, including those published by him in conjunction with his pupils, as well as the Monograph on the Elasmobranch Fishes. The last memoir in the volume, that on the Anatomy and Development of Peripatus Capensis, was published after his death, from his notes and drawings, with additions by Prof. Moseley and Mr Adam Sedgwick, who prepared the manuscript for publication. To the volume is prefixed an introductory biographical notice.

The second and third volumes are the two volumes of the Comparative Embryology reprinted from the original edition without alteration, save the correction of obvious misprints and omissions.

The fourth volume contains the plates illustrating the memoirs contained in Vol. i. We believe that we are consulting the convenience of readers in adopting this plan, rather than in distributing the plates among the memoirs to which they belong. To assist the reader the explanations of these plates have been given twice : at the end of the memoir to which they belong (in the case of the Monograph on Elasmobranch Fishes at the end of each separate chapter), and in the volume of plates.

All the figures of these plates had to be redrawn on the stone, and our best thanks are due to the Cambridge Scientific Instrument Company for the pains which they have taken in executing this work. We are also indebted to the Committee of Publication of the Zoological Society for the gift of electrotypes of the wood-cuts illustrating memoir no. xx. of Vol. i.

Several photographs of Balfour, taken at different times of his life, the last shortly before his death, are in the possession of his relatives and friends ; but these, in the opinion of many, leave much to be desired.

There 'is also a portrait of him in oils painted since his death by Mr John Collier, A.R.A., and Herr Hildebrand of Florence has executed a posthumous bust in bronze*. The portrait which forms the frontispiece of Vol. i. has been drawn on stone by Mr E. Wilson of the Cambridge Scientific Instrument Company, after the latest photograph. Should it fail, in the eyes of those who knew Balfour well, to have reproduced with complete success his features and expression, we would venture to ask them to bear in mind the acknowledged difficulties of posthumous portraiture.

  • In possession of the family. Copies also exist in the Library of

Trinity College, and in the Morphological Laboratory, at Cambridge.

Table of Contents

  1. On some points in the Geology of the East Lothian Coast. By G. W. and F. M. BALFOUR 1873
  2. The development and growth of the layers of the blastoderm. With Plate i 29
  3. On the disappearance of the Primitive Groove in the Embryo Chick With Plate i 41
  4. The development of the blood-vessels of the Chick. With Plate 2 47 1874
  5. A preliminary account of the development of the Elasmobranch FishesWith Plates 3 and 4 1875
  6. A comparison of the early stages in the development of Vertebrates With Plate 5 112
  7. On the origin and history of the urinogenital organs of Vertebrates 135
  8. On the development of the spinal nerves in Elasmobranch Fishes With Plates 22 and 23 1876
  9. On the spinal nerves of Amphioxus 1876-78
  10. A Monograph on the development of Elasmobranch Fishes. With Plates 1878
  11. On the phenomena accompanying the maturation and impregnation of the ovum
  12. On the structure and development of the vertebrate ovary With Plates 24, 25, 26 1879
  13. On the existence of a Head-kidney in the Embryo Chick, and on certain points in the development of the Müllerian duct By F. M. Balfour And A. Sedgwick. With Plates 27 and 28
  14. On the early development of the Lacertilia, together with some observations on the nature and relations of the primitive Streak With Plate 29
  15. On certain points in the Anatomy of Peripatus Capensis
  16. On the morphology and systematic position of the Spongida 1880
  17. Notes on the development of the Araneina With Plates 30, 31, 32
  18. On the spinal nerves of Amphioxus
  19. Address to the Department of Anatomy and Physiology of the British Association for the Advancement of Science 1881
  20. On the development of the skeleton of the paired fins of Elasmobranchii, considered in relation to its bearings on the nature of the limbs of the Vertebrata With Plate 33
  21. On the evolution of the Placenta, and on the possibility of employing the characters of the Placenta in the classification of the Mammalia 1882
  22. On the structure and development of Lepidosteus By F. M. BALFOUR and W. N. PARKER. With Plates 34-42
  23. On the nature of the organ in Adult Teleosteans and Ganoids which is usually regarded as the Head-kidney or Pronephros
  24. A renewed study of the germinal layers of the Chick By F. M. BALFOUR and F. DEIGHTON. With Plates 43,44,45
  25. The Anatomy and Development of Peripatus Capensis Edited by H. N. MoSELEYand A. SEDGWICK. With Plates 46-53 POSTHUMOUS, 1883


Francis Maitland Balfour, the sixth child and third son of James Maitland Balfour of Whittinghame, East Lothian, and Lady Blanche, daughter of the second Marquis of Salisbury, was born at Edinburgh, during a temporary stay of his parents there, on the loth November, 1851. He can hardly be said to have known his father, who died of consumption in 1856, at the early age of thirty-six, and who spent the greater part of the last two years of his life at Madeira, separated from the younger children who remained at home. He fancied at one time that he had inherited his father's constitution ; and this idea seems to have spurred him on to achieve early what he had to do. But, though there was a period soon after he went to College, during which he seemed delicate, and the state of his health caused considerable anxiety to his friends, he eventually became fairly robust, and that in spite of labours which greatly taxed his strength.

The early years of his life were spent chiefly at Whittinghame under the loving care of his mother. She made it a point to attempt to cultivate in all her children some taste for natural science, especially for natural history, and in this she was greatly helped by the boys' tutor, Mr J. W. Kitto. They were encouraged to make collections and to form a museum, and the fossils found in the gravel spread in front of the house served as the nucleus of a geological series. Frank soon became greatly interested in these things, and indeed they may be said to have formed the beginnings of his scientific career. At all events there was thus awakened in him a love for geology, which science continued to be his favorite study all through his boyhood, and interested him to the last. He was most assiduous in searching for fossils in the gravel and elsewhere, and so great was his love for his collections that while as yet quite a little boy the most delightful birthday present he could think of was a box with trays and divisions to hold his fossils and specimens. His mother, thinking that his fondness for fossils was a passing fancy and that he might soon regret the purchase of the box, purposely delayed the present. But he remained constant to his wish and in time received his box. He must at this time have been about seven or eight years old. In the children's museum, which has been preserved, there are specimens labelled with his childish round-hand, such as a piece of stone with the label " marks of some shels ;" and his sister Alice, who was at that time his chief companion, remembers discussing with him one day after the nursery dinner, when he was about nine years old, whether it were better to be a geologist or a naturalist, he deciding for the former on the ground that it was better to do one thing thoroughly than to attempt many branches of science and do them imperfectly.

Besides fossils, he collected not only butterflies, as do most boys at some time or other, but also birds ; and he with his sister Alice, being instructed in the art of preparing and preserving skins, succeeded in making a very considerable collection. He thus acquired before long not only a very large but a very exact knowledge of British birds.

In the more ordinary work of the school-room he was somewhat backward. This may have been partly due to the great difficulty he had in learning to write, for he was not only lefthanded but, in his early years, singularly inapt in acquiring particular muscular movements, learning to dance being a great trouble to him. Probably however the chief reason was that he failed to find any interest in the ordinary school studies. He fancied that the family thought him stupid, but this does not appear to have been the case.

In character he was at this time quick tempered, sometimes even violent, and the energy which he shewed in after life even thus early manifested itself as perseverance, which, when he was crossed, often took on the form of obstinacy, causing at times no little trouble to his nurses and tutors. But he was at the same time warm-hearted and affectionate ; full of strong impulses, he disliked heartily and loved much, and in his affections was wonderfully unselfish, wholly forgetting himself in his thought for others, and ready to do things which he disliked to please those whom he loved. Though, as we have said, somewhat clumsy, he was nevertheless active and courageous ; in learning to ride he shewed no signs of fear, and boldly put his pony to every jump which was practicable.

In 1 86 1 he was sent to the Rev. C. G. Chittenden's preparatory school at Hoddesden in Hertfordshire, and here the qualities which had been already visible at home became still more obvious. He found difficulty not only in writing but also in spelling, and in the ordinary school-work he took but little interest and made but little progress.

In 1865 he was moved to Harrow and placed in the house of the Rev. F. Rendall. Here, as at Hoddesden, he did not shew any great ability in the ordinary school studies, though as he grew older his progress became more marked. But happily he found at Harrow an opportunity for cultivating that love of scientific studies which was yearly growing stronger in him. Under the care of one of the Masters, Mr G. Griffith, the boys at Harrow were even then taught the elements of natural science. The lessons were at that time, so to speak, extraacademical, carried on out of school hours ; nevertheless, many of the boys worked at them with diligence and even enthusiasm, and among these Balfour became conspicuous, not only by his zeal but by his ability. Griffith was soon able to recognize the power of his new pupil, and thus early began to see that the pale, earnest, somewhat clumsy-handed lad, though he gave no promise of being a scholar in the narrower sense of the word, had in him the makings of a man of science. Griffith chiefly confined his teaching to elementary physics and chemistry with some little geology, but he also encouraged natural history studies and began the formation of a museum of comparative anatomy. Balfour soon began to be very zealous in dissecting animals, and was especially delighted when the Rev. A. C. Eaton, the well-known entomologist, on a visit to Harrow, initiated Griffith's pupils in the art of dissecting under water. The dissection of a caterpillar in this way was probably an epoch in Balfour's life. Up to that time his rough examination of such bodies had revealed to him nothing more than what in school-boy language he spoke of as " squash ;" but when under Eaton's deft hands the intricate organs of the larval Arthropod floated out under water and displayed themselves as a labyrinth of threads and sheets of silvery whiteness a new world of observation opened itself up to Balfour, and we may probably date from this the beginning of his exact morphological knowledge.

While thus learning the art of observing, he was at the same time developing his power of thinking. He was by nature fond of argument, and defended with earnestness any opinions which he had been led to adopt. He was very active in the Harrow Scientific Society, reading papers, taking part in the discussions, and exhibiting specimens. He gained in 1867 a prize for an essay on coal, and when, in 1868, Mr Leaf offered a prize (a microscope) "for the best account of some locality visited by the writer during the Easter Holidays," two essays sent in, one by Balfour, the other by his close friend, Mr Arthur Evans, since well known for his researches in Illyria, were found to be of such unusual merit that Prof. Huxley was specially requested to adjudicate between them. He judged them to be of equal merit, and a prize was given to each. The subject of Balfour's essay was " The Geology and Natural History of East Lothian." When biological subjects were discussed at the Scientific Society, Balfour appears to have spoken as a most uncompromising opponent of the views of Mr Charles Darwin, little thinking that in after life his chief work would be to develope and illustrate the doctrine of evolution.

The years at Harrow passed quickly away, Balfour making fair, but perhaps not more than fair, progress in the ordinary school learning. In due course however he reached the upper sixth form, and in his last year, became a monitor. At the same time his exact scientific knowledge was rapidly increasing. Geology still continued to be his favorite study, and in this he made no mean progress. During his last years at Harrow he and his brother Gerald worked out together some views concerning the geology of their native county. These views they ultimately embodied in a paper, which was published in their joint names in the Geological Magazine for 1872, under the title of "Some Points in the Geology of the East Lothian Coast," and which was in itself a work of considerable promise. Geology however was beginning to find a rival in natural history. Much of his holiday time was now spent in dredging for marine animals along the coast off Dunbar. Each specimen thus obtained was carefully determined and exact records were kept of the various ' finds,' so that the dredgings (which were zealously continued after he had left Harrow and gone to Cambridge) really constituted a serious study of the fauna of this part of the coast. They also enabled him to make a not inconsiderable collection of shells, in the arrangement of which he was assisted by his sister Evelyn, of Crustacea and of other animals.

Both to the masters and to his schoolfellows he became known as a boy of great force of character. Among the latter his scrupulous and unwavering conscientiousness made him less popular perhaps than might have been expected from his bright kindly manner and his unselfish warmheartedness. In the incidents of school life a too strict conscience is often an inconvenience, and the sternness and energy with which Balfour denounced acts of meanness and falsehood were thought by some to be unnecessarily great. He thus came to be feared rather than liked by many, and comparatively few grew to be sufficiently intimate with him to appreciate the warmth of his affections and the charm of his playful moments.

At the Easter of 1870 he passed the entrance examination at Trinity College, Cambridge, and entered into residence in the following October. His college tutor was Mr J. Prior, but he was from the first assisted and guided in his studies by his friend, Mr Marlborough Pryor, an old Harrow boy, who in the same October had been, on account of his distinction in Natural Science, elected a Fellow of the College, in accordance with certain new regulations which then came into action for the first time, and which provided that every three years one of the College Fellowships should be awarded for excellence in some branch or branches of Natural Science, as distinguished from mathematics, pure or mixed. During the whole of that year and part of the next Mr Marlborough Pryor remained in residence, and his influence in wisely directing Balfour's studies had a most beneficial effect on the latter's progress.

During his first term Balfour was occupied in preparation for the Previous Examination ; and this he successfully passed at Christmas. After that he devoted himself entirely to Natural Science, attending lectures on several branches. During the Lent term he was a very diligent hearer of the lectures on Physiology which I was then giving as Trinity Praelector, having been appointed to that post in the same October that Balfour came into residence. At this time he was not very strong, and I remember very well noticing among my scanty audience, a pale retiring student, whose mind seemed at times divided between a desire to hear the lecture and a feeling that his frequent coughing was growing an annoyance to myself and the class. This delicate-looking student, I soon learnt, was named Balfour, and when the Rev. Coutts Trotter, Mr Pryor and myself came to examine the candidates for the Natural Science Scholarships which were awarded at Easter, we had no difficulty in giving the first place to him. In point of knowledge, and especially in the thoughtfulness and exactitude displayed in his papers and work, he was very clearly ahead of his competitors.

During the succeeding Easter term and the following winter he appears to have studied physics, chemistry, geology and comparative anatomy, both under Mr Marlborough Pryor and by means of lectures. He also continued to attend my lectures, but though I gradually got to know him more and more we did not become intimate until the Lent term of 1872. He had been very much interested in some lectures on embryology which I had given, and, since Marlborough Pryor had left or was about to leave Cambridge, he soon began to consult me a good deal about his studies. He commenced practical histological and embryological work under me, and I remember very vividly that one day when we were making a little excursion in search of nests and eggs of the stickleback in order that he might study the embryology of fishes, he definitely asked my opinion as to whether he might take up a scientific career with a fair chance of success. I had by this time formed a very high opinion of his abilities, and learning then for the first time that he had an income independent of his own exertions, my answer was very decidedly a positive one. Soon after, feeling more and more impressed with his power and increasingly satisfied both with his progress in biological studies and his sound general knowledge of other sciences, anxious also, it may be, at the same time that as much original inquiry as possible should be carried on at Cambridge in my department, I either suggested to him or acquiesced in his own suggestion that he should at once set to work on some distinct research ; and as far as I remember the task which I first proposed to him was an investigation of the layers of the blastoderm in the chick. It must have been about the same time that I proposed to him to join me in preparing for publication a small work on Embryology, the materials for this I had ready to hand in a rough form as lectures which. I had previously given. To this proposal he enthusiastically assented, and while the lighter task of writing what was to be written fell to me, he undertook to work over as far as was possible the many undetermined points and unsatisfactory statements across which we were continually coming.

During his two years at College his health had improved ; though still hardly robust and always in danger of overworking himself, he obviously grew stronger. He rejoiced exceedingly in his work, never tiring of it, and was also making his worth felt among his fellow students, and especially perhaps among those of his own college whose studies did not lie in the same direction as his own. At this time he must have been altogether happy, but a sorrow now came upon him. His mother, to whom he was passionately attached, and to whose judicious care in his early days not only the right development of his strong character but even his scientific leanings were due, had for some time past been failing in health, though her condition caused no immediate alarm. In May 1872, however, she died quite suddenly from unsuspected heart disease. Her loss was a great blow to him, and for some time afterward I feared his health would give way ; but he bore his grief quietly and manfully and threw himself with even increased vigour into his work.

During the academic session of 1872 3, he continued steadily at work at his investigations, and soon began to make rapid progress. At the beginning he had complained to me about what he considered his natural clumsiness, and expressed a fear that he should never be able to make satisfactory microscopic sections ; as to his being able to make drawings of his dissections and microscopical preparations, he looked upon that at first as wholly impossible. I need hardly say that in time he acquired great skill in the details of microscopical technique, and that his drawings, if wanting in so-called artistic finish, were always singularly true and instructive. While thus struggling with the details which I could teach him, he soon began to manifest qualities which no teacher could give him. I remember calling his attention to Dursy's paper on the Primitive Streak, and suggesting' that he should work the matter over, since if such a structure really existed, it must, most probably, have great morphological significance. I am free to. confess that I myself rather doubted the matter, and a weaker student might have been influenced by my preconceptions. Balfour, however, thus early had the power of seeing what existed and of refusing to see what did not exist. He was soon able to convince me that Dursy's streak was a reality, and the complete working out of its significance occupied his thoughts to the end of his days.

The results of these early studies were made known in three papers which appeared in the Quarterly Journal of Microscopical Science for July 1873, and will be found in the beginning of this volume. The summer and autumn of that year were spent partly in a visit to Finland, in company with his friend and old schoolfellow Mr Arthur Evans, and partly in formal preparation for the approaching Tripos examination. Into this preparation Balfour threw himself with characteristic energy, and fully justified my having encouraged his spending so much of the preceding time in original research, not only by the rapidity with which he accumulated the stock of knowledge of various kinds necessary for the examination but also by the manner in which he acquitted himself at the trial itself. At that time the position of the candidates in the Natural Sciences Tripos was determined by the total number of marks, and Balfour was placed second, the first place being gained by H. Newell Martin of Christ's College, now Professor at Baltimore, U.S.A. In the examination, in which I took part, Balfour did not write much, and he had not yet learnt the art of putting his statements In the best possible form ; he won his position chiefly by the firm thought and clear insight which was present in almost all his answers.

The examination was over in the early days of Dec. 1873 and Balfour was now free to devote himself wholly to his original work. Happily, the University had not long before secured the use of two of the tables at the then recently founded Stazione Zoologica at Naples. And upon the nomination of the University, Balfour, about Christmas, started for Naples in company with his friend Mr A. G. Dew-Smith, also of Trinity College. The latter was about to carry on some physiological observations ; Balfour had set himself to work out as completely as he could the embryology of Elasmobranch fishes, about which little was at that time known, but which, from the striking characters of the adult animals could not help proving of interest and importance.

From his arrival there at Christmas 1873 until he left in June 1874, he worked assiduously, and with such success, that as the result of the half-year's work he had made a whole series of observations of the greatest importance. Of these perhaps the most striking were those on the development of the urogenital organs, on the neurenteric canal, on the development of the spinal nerves, on the formation of the layers and on the phenomena of segmentation, including a history of the behaviour of nuclei in cell division. He returned home laden with facts and views both novel and destined to influence largely the progress of embryology.

In August of the same year he attended the meeting of the British Association for the Advancement of Science at Belfast ; and the account he then gave of his researches formed one of the most important incidents at the Biological Section on that occasion.

In the September of that year the triennial fellowship for Natural Science was to be awarded at Trinity College, and Balfour naturally was a candidate. The election was, according to the regulations, to be determined partly by the result of an examination in various branches of science, and partly by such evidence of ability and promise as might be afforded by original work, published or in manuscript. He spent the remainder of the autumn in preparation for this examination. But when the examination was concluded it was found that in his written answers he had not been very successful ; he had not even acquitted himself so well as in the Tripos of the year before, and had the election been determined by the results of the examination alone, the examiners would have been led to choose the gentleman who was Balfour's only competitor. The original work however which Balfour sent in, including a preliminary account of the discoveries made at Naples, was obviously of so high a merit and was spoken of in such enthusiastic terms by the External Referee Prof. Huxley, that the examiners did not hesitate for a moment to neglect altogether the formal written answers (and indeed the papers of questions were only introduced as a safeguard, or as a resource in case evidence of original power should be wanted) and unanimously recommended him for election. Accordingly he was elected Fellow in the early days of October.

Almost immediately after, the little book on Embryology appeared, on which he and I had been at work, he doing his share even while his hands and mind were full of the Elasmobranch inquiry. The title-page was kept back some little time in order that his name might appear on it with the addition of Fellow of Trinity, a title of which he was then, and indeed always continued to be, proud. He also published in the October number of the Quarterly Journal of Microscopical Science a preliminary account of his Elasmobranch researches.

He and his friends thought that after these almost incessant labours, and the excitement necessarily contingent upon the fellowship election, he needed rest and change. Accordingly on the i /th of October he started with his friend Marlborough Pryor on a voyage to the west coast of South America. They travelled thither by the Isthmus of Panama, visited Peru and Chili, and returned home along the usual route by the Horn ; reaching England some time in Feb. 1875.

Refreshed by this holiday, he now felt anxious to complete as far as possible his Elasmobranch work, and very soon after his return home, in fact in March, made his way again to Naples, where he remained till the hot weather set in in May.

On his return to Cambridge, he still continued working on the Elasmobranchs, receiving material partly from Naples, partly from the Brighton Aquarium, the then director of which, Mr Henry Lee, spared no pains to provide him both with embryo and adult fishes. While at Naples, he communicated to the Philosophical Society at Cambridge a remarkable paper on "The Early Stages of Vertebrates," which was published in full in the Quarterly Journal of Microscopical Science, July, 1875; he also sent me a paper on "The Development of the Spinal Nerves", which I communicated to the Royal Society, and which was subsequently published in the Philosophical Transactions of 1876. He further wrote in the course of the summer and published in the Journal of Anatomy and Physiology \\\ October, 1875, a detailed account of his "Observations and Views on the Development of the Urogenital Organs."

Some time in August of the same year he started in company with Mr Arthur Evans and Mr J. F. Bullar for a second trip to Finland, the travellers on this occasion making their way into regions very seldom visited, and having to subsist largely on the preserved provisions which they carried with them, and on the produce of their rods and guns. From a rough diary which Balfour kept during this trip it would appear that while enjoying heartily the fun of the rough travelling, he occupied himself continually with observations on the geology and physical phenomena of the country, as well as on the manners, antiquities, and even language of the people. It was one of his characteristic traits, a mark of the truly scientific bent of his mind, of his having, as Dohrn soon after Balfour's first arrival at Naples said, ' a real scientific head,' that every thing around him wherever he was, incited him to careful exact observation, and stimulated him to thought.

In the early part of the Long Vacation of the same year he had made his first essay, in lecturing, having given a short course on Embryology in a room at the New Museums, which I then occupied as a laboratory. Though he afterwards learnt to lecture with great clearness he was not by nature a fluent speaker, and on this occasion he was exceedingly nervous. But those who listened to him soon forgot these small defects as they began to perceive the knowledge and power which lay in their new teacher.

Encouraged by the result of this experiment, he threw himself, in spite of the heavy work which the Klasmobranch investigation was entailing, with great zeal into an arrangement which Prof. Newton, Mr J. W. Clark and myself had in course of the summer brought about, that he and Mr A. Milnes Marshall, since Professor at Owens College, Manchester, should between them give a course on Animal Morphology, with practical instruction, Prof. Newton giving up a room in the New Museums for the purpose.

In the following October (1*87 5) upon Balfour's return from Finland, these lectures were accordingly begun and carried on by the two lecturers during the Michaelmas and Lent Terms. The number of students attending this first course, conducted on a novel plan, was, as might be expected, small, but the Lent Term did not come to an end before an enthusiasm for morphological studies had been kindled in the members of the class.

The ensuing Easter term (1876) was spent by Balfour at Naples, in order that he might carry on towards completion his Elasmobranch work. He had by this time determined to write as complete a monograph as he could of the development of these fishes, proposing to publish it in instalments in the Journal of Anatomy and Physiology, and subsequently to gather together the several papers into one volume. The first of these papers, dealing with the ovum, appeared in Jan. 1876; most of the numbers of the Journal during that and the succeeding year contained further portions ; but the complete monograph did not leave the publisher's hands until 1878.

He returned to England with his pupil and friend Mr J. F. Bullar some time in the summer ; on their way home they passed through Switzerland, and it was during the few days which he then spent in sight of the snow-clad hills that the beginnings of a desire for that Alpine climbing, which was destined to be so disastrous, seem to have been kindled in him.

In October, 1876, he resumed the lectures on Morphology, taking the whole course himself, his colleague, Mr Marshall, having meanwhile left Cambridge. Indeed, from this time onward, he may be said to have made these lectures, in a certain sense, the chief business of his life. He lectured all three terms, devoting the Michaelmas and Lent terms to a systematic course of Animal Morphology, and the Easter term to a more elementary course of Embryology. These lectures were given under the auspices of Prof. Newton ; but Balfour's position was before long confirmed by his being made a Lecturer of Trinity College, the lectures which he gave at the New Museums, and which were open to all students of the University, being accepted in a liberal spirit by the College as equivalent to College Lectures. He very soon found it desirable to divide the morphological course into an elementary and an advanced course, and to increase the number of his lectures from three to four a week. Each lecture was followed by practical work, the students dissecting and examining microscopically, an animal or some animals chosen as types to illustrate the subject-matter of the lecture ; and although Balfour had the assistance at first of one 1 , and ultimately of several demonstrators, he himself put his hand to the plough, and after the lecture always spent some time in the laboratory among his pupils. Had Balfour been only an ordinary man, the zeal and energy which he threw into his lectures, and into the supervision of the practical work, added to the almost brotherly interest which he took in the individual development of every one of the pupils who shewed any love whatever for the subject, would have made him a most successful teacher. But his talents and powers were such as could not be hid even from beginners. His extensive and exact knowledge, the clearness with which in spite of, or shall I not rather say, by help of a certain want of fluency, he explained difficult and abstruse matters, the trenchant way in which he laybare specious fallacies, and the presence in almost his every word of that power which belongs only to the man who has thought out for himself everything which he says, these things aroused and indeed could hardly fail to arouse in his hearers feelings which, except in the case of the very dullest, grew to be those of enthusiasm. His class, at first slowly, but afterwards more rapidly, increased in numbers, and, what is of more importance, grew in quality. The room allotted to him soon became far too small and steps were taken to provide for him, for myself, whose wants were also urgent, and for the biological studies generally, adequate accommodation ; but it was not until Oct. 1877 that we were able to take possession of the new quarters.

  • 1 His first Demonstrator up to Christinas 1877, was Mr J. F. Bullar. In Jan.

1878, Mr Adam Sedgwick took the post of Senior Demonstrator, and held it until Balfour's death.

Even this new accommodation soon became insufficient, and in the spring of 1882 a new morphological laboratory was commenced in accordance with plans suggested by himself. He was to have occupied them in the October term, 1883, but did not live to see them finished.

As might have been expected from his own career, he regarded the mere teaching of what is known as a very small part of his duties as Lecturer ; and as soon as any of his pupils became sufficiently advanced, he urged or rather led them to undertake original investigations ; and he had the satisfaction before his death of seeing the researches of his pupils (such as those by Messrs. Bullar, Sedgwick, Mitzikuri, Haddon, Scott, Osborne, Caldwell, Heape, Weldon, Parker, Deighton and others) carried to a successful end. In each of these inquiries he himself took part, sometimes a large part, generally suggesting the problem to be solved, indicating the methods, and keeping a close watch over the whole progress of the study. Hence in many cases the published account bears his name as well as that of the pupil.

In the year 1878 his Monograph on Elasmobranch Fishes was published as a complete volume, and in the same year he received the honour of being elected a Fellow of the Royal Society, a distinction which now-a-days does not often fall to one so young. No sooner was the Monograph completed than in spite of the labours which his lectures entailed, he set himself to the great task of writing a complete treatise on Comparative Embryology. This not only laid upon him the heavy burden of gathering together the observations of others, enormous in number and continually increasing, scattered through many journals and books, and recorded in many different languages, as well as of putting them in orderly array, and of winnowing out the grain from the chaff (though his critical spirit found some relief in the latter task), but also caused him much labour, inasmuch as at almost every turn new problems suggested themselves, and demanded inquiry before he could bring his mind to writing about them. This desire to see his way straight before him, pursued him from page to page, and while it has resulted in giving the book an almost priceless value, made the writing of it a work of vast .labour. Many of the ideas thus originated served as the bases of inquiries worked out by himself or his pupils, and published in the form of separate papers, but still more perhaps never appeared either in the book or elsewhere and were carried with him undeveloped and unrecorded to the grave.

The preparation of this work occupied the best part of his time for the next three years, the first volume appearing in 1880, the second in 1881.

In the autumn of 1880, he attended the Meeting at Swansea of the British Association for the Advancement of Science, having been appointed Vice-President of the Biological Section with charge of the Department of Anatomy and Physiology. At the Meetings of the Association, especially of late years, much, perhaps too much, is expected in the direction of explaining the new results of science in a manner interesting to the unlearned. Popular expositions were never very congenial to Balfour, his mind was too much occupied with the anxiety of problems yet to be solved ; he was therefore not wholly at his ease, in his position on this occasion. Yet his introductory address, though not of a nature to interest a large mixed audience, was a luminous, brief exposition of the modern development and aims of embryological investigation.

During these years of travail with the Comparative Embryology the amount of work which he got through was a marvel to his friends, for besides his lectures, and the researches, and the writing of the book, new labours were demanded of him by the University for which he was already doing so much. Men at Cambridge, and indeed elsewhere as well, soon began to find out that the same clear insight which was solving biological problems could be used to settle knotty questions of policy and business. Moreover he united in a remarkable manner, the power of boldly and firmly asserting and maintaining his own views, with a frank courteousness which went far to disarm opponents. Accordingly he found himself before long a member of various Syndicates, and indeed a very great deal of his time was thus occupied, especially with the Museums and Library Syndicates, in both of which he took the liveliest interest. Besides these University duties his time and energy were also at the service of his College. In the preparation of the New Statutes, with which about this time the College was much occupied, the Junior Fellows of the College took a conspicuous share, and among these Junior Fellows Balfour was perhaps the most active ; indeed he was their leader, and he threw himself into the investigation of the bearings and probable results of this and that proposed new statute with as much zeal as if he were attacking some morphological problem.

While he was in the midst of these various labours, his friends, often feared for his strength, for though gradually improving in health after his first year at Cambridge, he was not robust, and from time to time he seemed on the point of breaking down. Still, hard as he was working, he was in reality wisely careful of himself, and as he grew older, paid more and more attention to his health, daily taking exercise in the form either of bicycle rides or of lawn-tennis. Moreover he continued to spend some part of his vacations in travel. Combining business with pleasure, he made frequent visits to Germany and France, and especially to Naples. The Christmas of 1876 7 he spent in Greece, that of 1878 9 at Ragusa, where his old school-fellow and friend Mr Arthur Evans was at that time residing, and the appointment of his friend Kleinenberg to a Professorship at Messina led to a journey there. Early in the long vacation of 1880, he went with his sister, Mrs H. Sidgwick, and her husband to Switzerland, and was joined there for a short time by his friend and pupil Adam Sedgwick. During this visit he took his first lessons in Alpine climbing, making several excursions, some of them difficult and dangerous ; and the love of mountaineering laid so firm a hold upon him, that he returned to Switzerland later on in the autumn of the same year, in company with his brother Gerald, and spent some weeks near Zermatt in systematic climbing, ascending, among other mountains, the Matterhorn and the Weisshorn. In the following summer, 1 88 1 , he and his brother Gerald again visited the Alps, dividing their time between the Chamonix district and the Bernese Oberland. On this occasion some of the excursions which they made were of extreme difficulty, and such as needed not only great presence of mind and bodily endurance, but also skilful and ready use of the limbs. As a climber indeed Balfour soon shewed himself fearless, indefatigable, and expert in all necessary movements as well as full of resources and expedients in the face of difficulties, so much so that he almost at once took rank among the foremost of distinguished mountaineers. In spite of his apparent clumsiness in some matters, he had even as a lad proved himself to be a bold and surefooted climber. Moreover he had been perhaps in a measure prepared for the difficulties of Alpine climbing by his experience in deerstalking. This sport he had keenly and successfully pursued for many years at his brother's place in Rosshire. When however about the year 1877, the question of physiological experiments on animals became largely discussed in public, he felt that to continue the pursuit of this or any other sport involving, for the sake of mere pleasure, the pain and death of animals, was inconsistent with the position which he had warmly taken up, as an advocate of the right to experiment on animals ; and he accordingly from that time onward wholly gave it up.

His fame as an investigator and teacher, and as a man of brilliant and powerful parts, was now being widely spread. Pupils came to him, not only from various parts of England, but from America, Australia and Japan. At the York Meeting of the British Association for the Advancement of Science, in August, 1 88 1, he was chosen as one of the General Secretaries. In April, 1881, the honorary degree of LL.D. was conferred upon him by the University of Glasgow, and in November of the same year the Royal Society gave him one of the Royal Medals in recognition of his embryological discoveries, and at the same time placed him on its Council.

At Cambridge he was chosen, in the autumn of 1880, President of the Philosophical Society, and in the December of that year a brilliant company were gathered together at the Annual Dinner to do honour to their new young President. Otherwise nothing as yet had been done for him in his own University in the way of recognition of his abilities and services ; and he still remained a Lecturer of Trinity College, giving lectures in a University building. An effort had been made by some of his friends to urge the University to take some step in this direction ; but it was thought at that time impossible to do anything. In 1 88 1 a great loss fell upon the sister University of Oxford in the death of Prof. George Rolleston ; and soon after very vigorous efforts were made to induce Balfour to become a candidate for the vacant chair. The prospect was in many ways a tempting one, and Balfour seeing no very clear way in the future for him at his own University, was at times inclined to offer himself, but eventually he decided to remain at Cambridge. Hardly had this temptation if we may so call it been overcome when a still greater one presented itself. Through the lamented death of Sir Wyville Thomson in the winter of 1 88 1 2, the chair of Natural History at Edinburgh, perhaps the richest and most conspicuous biological chair in the United Kingdom, became vacant. The post was in many ways one which Balfour would have liked to hold. The teaching duties were it is true laborious, but they had in the past been compressed into a short time, occupying only the summer session and leaving the rest of the year free, and it seemed probable that this arrangement might be continued with him. The large emolument would also have been grateful to him inasmuch as he would have felt able to devote the whole of it to scientific ends ; and the nearness to Whittinghame. his native place and brother's home, added to the attractions ; but what tempted him most was the position which it would have given him, and the opportunities it would have afforded, with the rich marine Fauna of the north-eastern coast close at hand, to develope a large school of Animal Morphology. The existing Professors at Edinburgh were most desirous that he should join them, and made every effort to induce him to come. On the part of the Crown, in whose hands the appointment lay, not only were distinct offers made to him, but he was repeatedly pressed to accept the post. Nor was it until after a considerable struggle that he finally refused, his love for his own University in the end overcoming the many inducements to leave ; he elected to stay where he was, trusting to the future opening up for him some suitable position. In this decision he was undoubtedly influenced by the consideration that Cambridge, besides being the centre of his old friendships, had become as it were a second home for his own family. By the appointment of Lord Rayleigh to the chair of Experimental Physics his sister Lady Rayleigh had become a resident, his sister Mrs Sidgwick had lived there now for some years, and his brother Gerald generally spent the summer there ; their presence made Cambridge doubly dear to him.

At the close of the Michaelmas term, with feelings of relief at having completed his Comparative Embryology, the preparation of the second volume of which had led to almost incessant labour during the preceding year, he started to spend the Christmas vacation with his friend Kleinenberg at Messina. Stopping at Naples on his way thither he found his pupil Caldwell, who had been sent to occupy the University table at the Stazione Zoologica, lying ill at Capri, with what proved to be typhoid fever. The patient was alone, without any friend to tend him, and his mother who had been sent for had not yet arrived. Accordingly Balfour (with the kindness all forgetful of himself which was his mark all his life through) stayed on his journey to nurse the sick man until the mother came. He then went on to Messina, and there seemed to be in good health, amusing himself with the ascent of Etna. Yet in January, soon after his return home, he complained of being unwell, and in due time distinct symptoms of typhoid fever made their appearance. The attack at first promised to be severe, but happily the crisis was soon safely passed and the convalescence was satisfactory.

While yet on his sick bed, a last attempt was made to induce him to accept the Edinburgh offer, and for the last time he refused. These repeated offers, and the fact that the dangers of his grave illness had led the University vividly to realize how much they would lose if Balfour were taken away from them, encouraged his friends to make a renewed effort to gain for him some adequate position in the University. This time the attempt was successful, and the authorities took a step, unusual but approved of by the whole body of resident members of the University ; they instituted a new Professorship of Animal Morphology, to be held by Balfour during his life or as long as he should desire, but to terminate at his death or resignation unless it should be otherwise desirable. Accordingly in May, 1882, he was admitted into the Professoriate as Professor of Animal Morphology.

During his illness his lectures had been carried on by his Demonstrator, Mr Adam Sedgwick, who continued to take his place during the remainder of that Lent Term and during the ensuing Easter Term. The spring Balfour spent partly in the Channel Islands with his sister Alice, partly in London with his eldest brother, but in the course of the Ea'ster Term returned to Cambridge and resumed his work though not his lectures. His recovery to health was steady and satisfactory, the only drawback being a swelling over the shin-bone of one leg, due to a blow on the rocks at Sark ; otherwise he was rapidly becoming strong. He himself felt convinced that a visit to the Alps, with some mountaineering of not too difficult a kind, would complete his restoration to health. In this view many of his friends coincided ; for the experience of former years had shewn them what a wonderfully beneficial effect the Alpine air and exercise had upon his health. He used to go away pale, thin and haggard, to return bronzed, clear, firm and almost stout ; nor was there anything in his condition which seemed to forbid his climbing, provided that he was cautious at the outset. Accordingly, early in June he left Cambridge for Switzerland, having long ago, during his illness in fact, engaged his old guide, Johann Petrus, whom he had first met in 1880, and who had always accompanied him in his expeditions since.

His first walking was in the Chamonix district ; and here he very soon found his strength and elasticity come back to him. Crossing over from Montanvert to Courmayeur, by the Col du Geant, he was attracted by the peak called the Aiguille Blanche de Peuteret, a virgin peak, the ascent of which had been before attempted but not accomplished. Consulting with Petrus he determined to try it, feeling that the fortnight, which by this time he had spent in climbing, had brought back to him his old vigour, and that his illness was already a thing of the past.

There is no reason to believe that he regarded the expedition as one of unusual peril ; and an incident which at the time of his death was thought by some to indicate this was in reality nothing more than a proof of his kindly foresight. The guide Petrus was burdened by a debt on his land amounting to about 150. In the previous year Balfour and his brother had come to know of this debt ; and, seeing that no Alpine ascent is free from danger, that on any expedition some accident might carry them off, had conceived the idea of making some provision for Petrus' family in case he might meet with sudden death in their service. This suggestion of the previous year Balfour carried out on this occasion, and sent home to his brother Gerald a cheque of 150 for this purpose. But the cheque was sent from Montanvert before he had even conceived the idea of ascending the Aiguille Blanche. It was not a provision for any specially dangerous ascent, and must be regarded as a measure prompted not by a sense of coming peril but rather by the donor's generous care for his servant.

On Tuesday afternoon, July 18, he and Petrus, with a porter to carry provisions and firing to their sleeping-place on the rocks, set out from Courmayeur, the porter returning the same night. They expected to get back to Courmayeur some time on the Thursday, but the day passed without their appearing. This did not cause any great anxiety because it was supposed that they might have found it more convenient to pass over to the Chamonix side than to return to Courmayeur. When on Friday however telegrams dispatched to Chamonix and Montanvert brought answers that nothing had been seen of them, it became evident that some accident had happened, and an exploring party set out for the hills. It was not until early on the Sunday morning that this search party found the bodies, both partly covered with snow, lying on the Glacier de Fresney, below the impassable icefall which separates the upper basin of the glacier from the lower portion, and at the foot of a couloir which descends by the side of the icefall. Their tracks were visible on the snow at the top of the couloir. Balfour's neck was broken, and his skull fractured in three places; Petrus' body was also fractured in many places. The exact manner of their death will never be known, but there can be no doubt that, in Balfour's case at all events, it was instantaneous, and those competent to form a judgment are of opinion that they were killed by a sudden fall through a comparatively small height, slipping on the rocks as they were descending by the side of the ice-fall, and not precipitated from the top of the couloir. There is moreover indirect evidence which renders it probable that in the fatal fall Petrus slipped first and carried Balfour with him. Whether they had reached the summit of the Aiguille and were returning home after a successful ascent or whether they were making their way back disheartened and wearied with failure, is not and perhaps never will be known. Since the provisions at the sleeping-place were untouched, the deaths probably took place on Wednesday the i pth. The bringing down the bodies proved to be a task of extreme difficulty, and it was not till Wednesday the 26th that the remains reached Courmayeur, where M. Bertolini, the master of the hotel, and indeed everyone, not least the officers of a small body of Italian troops stationed there, shewed the greatest kindness and sympathy to Balfour's brothers, Gerald and Eustace, who hastened to the spot as soon as the news of the terrible disaster was telegraphed home. Mr Walter Leaf also and Mr Conway, friends of Balfour, the former a very old one, who had made their way to Courmayeur from other parts of Switzerland as soon as they heard of the accident, rendered great assistance. The body was embalmed, brought to England, and buried at Whittinghame on Saturday, Aug. 5, the Fellows of Trinity College holding a service in the College Chapel at the same time.

In person he was tall, being fully six feet in height, well built though somewhat spare. A broad forehead overhanging deeply set dark brown eyes whose light shining from beneath strongly marked eye-brows told all the changes of his moods, slightly prominent cheek-bones, a pale skin, at times inclined to be even sallow, dark brown hair, allowed to grow on the face only as a small moustache, and slight whiskers, made up a countenance which bespoke at once strength of character and delicacy of constitution. It was an open countenance, hiding nothing, giving sign at once, both when his body was weary or weak, and when his mind was gladdened, angered or annoyed.

The record of some of his thoughts and work, all that he had given to the world will be found in the following pages. But who can tell the ideas which had passed into his quick brain, but which as yet were known only to himself, of which he had given no sign up to that sad day on which he made the fatal climb? And who can say whither he might not have reached had he lived, and his bright young life ripened as years went on ? This is not the place to attempt any judgment of his work : that may be left to other times, and to other hands; but it may be fitting to place here on record a letter which shews how much the greatest naturalist of this age appreciated his younger brother. Among Balfour's papers was found a letter from Charles Darwin, acknowledging the receipt of Vol. II. of the Comparative Embryology in the following words :

"July 6, 1881.



I thank you heartily for the present of your grand book, and I congratulate you on its completion. Although I read almost all of Vol. I. I do not feel that I am worthy of your present, unless indeed the fullest conviction that it is a memorable work makes me worthy to receive it.

  • * * * *

Once again accept my thanks, for I am proud to receive a book from you, who, I know, will some day be the chief of the English Biologists.

Believe me,

Yours sincerely,


The loss of him was a manifold loss. He is mourned, and will long be mourned, for many reasons. Some miss only the brilliant investigator ; others feel that their powerful and sympathetic teacher is gone ; some look back on his memory and grieve for the charming companion whose kindly courtesy and bright wit made the hours fly swiftly and pleasantly along ; and to yet others is left an aching void when they remember that they can never again lean on the friend whose judgment seemed never to fail and whose warm-hearted affection was a constant help. And to some he was all of these. At the news of his death the same lines came to the lips of all of us, so fittingly did Milton's words seem to speak our loss and grief

"For Lycidas is dead, dead ere his prime, Young Lycidas, and hath not left his peer."


The Works of Francis Balfour 1-1