John Goodsir FRS (1814–1867): Pioneer of cytology and microbiology

Introduction

John Goodsir^1–3 was a scientist who stimulated research throughout Britain. His interest in nature, encouraged by a childhood spent close to the sea, led to a fascination with every form of animal life. The compound microscope was emerging as an important research tool ⁴ and Goodsir’s work on cytology was a factor leading Rudolph Virchow to his epoch-making Cellular Pathology, ⁵ dedicated to Goodsir.

Early years

Goodsir’s forbears came to Fife, Scotland as Huguenot asylum-seekers. ⁶ John Goodsir was born on 20 March 1814, his father a medical practitioner in Anstruther. His younger brother, Joseph Taylor, born in 1816, entered the church in Edinburgh. A third brother, Henry Duncan Spens (Harry), perished during the 1845 Franklin expedition. ⁷ The fourth brother, Robert Anstruther, born in 1824, qualified in Medicine and sailed twice to the Arctic searching for his brother Harry. The youngest son, Archibald Johnstone, born in 1826 at Largo, qualified as a Member of the Royal College of Surgeons of England, the RCSEng, and served as a surgeon in the Fifeshire regiment of militia.

John Goodsir, a tall, gaunt figure, ⁸ attended school at Anstruther. He was taught drawing by his mother who encouraged his interest in the marine life he found as he searched the Largo coastal waters. One day he encountered Professor James Syme, invited by Goodsir’s father to examine a patient. Goodsir impressed the eminent surgeon by explaining the structure of calculi found on the beach.

In 1827, John Goodsir began studies at St Andrews University that included classics and mathematics, a course that ended in 1830. His ambition was to become a surgeon. Goodsir’s father hoped that his son would qualify from the Royal College of Surgeons of Edinburgh, the RCSEd, and work for James Syme or Robert Liston. Meanwhile Goodsir was offered an apprenticeship with the Edinburgh dentist Robert Naysmith, allowing him to matriculate at the University Medical School and to attend classes both there and at the RCSEd. He began to collect human teeth and discovered that deciduous teeth were not the ‘parents’ of the permanent teeth: they developed independently. ⁹ In 1840, he spoke to the British Association for the Advancement of Science, the BA, on ‘Dentition in the ruminants’, ¹⁰ assisted by the Professor of Natural History, Robert Jameson, who lent him an Ehrenburg microscope.

Among Goodsir’s Edinburgh teachers were Ferguson (Anatomy) and Syme (Clinical Surgery). Robert Christison now taught Materia Medica and Therapeutics, Hope (Chemistry) and Graham (Botany). Meanwhile Robert Knox, ¹¹ teaching in the Extramural School of Medicine, was reviving Edinburgh’s fame. The University and RCSEd museums stimulated Goodsir’s interest in human as well as zoological forms while the classrooms brought out his talent for dissection and drawing. Knox proposed Goodsir as Vice-President of the Anatomical and Physiological Society, the APS, and subsequently invited him to assist in the translation of Blainville’s De l’Organisation des Animaux. ¹²

By 1833, Goodsir had come to find dentistry wearisome. Naysmith agreed to cancel his indenture but employed him as a locum until 1835, the year in which Goodsir qualified as a Licentiate of the RCSEd. He then joined his father’s medical practice in Anstruther, giving him the continued opportunity of searching the coast for all forms of life. The specimens he accumulated initiated the collection that he developed when he became a museum conservator.

In Knox’s rooms, Goodsir formed a close friendship with the young Edward Forbes ¹³ from the Isle of Man. Goodsir concentrated on Anatomy, Pathology and Morphology while Forbes chose to study European Natural History and shortly before his premature death in 1854, succeeded Robert Jameson as Edinburgh University Regius Professor in this subject. Goodsir showed Forbes how to dissect a land snail, Clausilia bidentata, a two-toothed snail of the phylum Mollusca, which he had found on Arthur’s Seat. Forbes took Goodsir to a meeting of the Royal Physical Society, the RPS, in 1834 where Goodsir was obliged to speak in public for the first time.

Goodsir, Forbes and George Day secured an Edinburgh flat at 21 Lothian Street for an annual rent of £17. Day would later be Chandos Professor of Anatomy and Medicine of St Andrews University. It was the meeting place for the scientists, writers and artists who constituted a Universal Brotherhood of Friends of Truth and where they enjoyed social occasions that attracted musicians like Theodore Martin and poets like James Ballantine and John Stuart Blackie. Visitors included Robert Knox, Samuel Brown, John Balfour the botanist and, almost certainly, John Hughes Bennett, Professor of the Institutes of Medicine as Physiology was termed.^14,15 In the flat the Friends kept a monkey, a dog, guinea pigs and a tortoise together with caged birds that included an eagle. In the attic they made an aquarium for frogs, fish and molluscs. The shelves were repositories for fossils, geological and archaeological specimens but also a place for Forbes’ ‘objets d’art’. Goodsir’s brother Harry named the flat The Barracks. Forbes preferred Our Palace at Edinburgh.

Museum conservator

In April 1839, Knox invited Goodsir to join him as a lecturer. Goodsir was deterred by his negotiations with the Faculty of Medicine regarding the Conservatorship of the University Anatomy collection. By May of that year, money for the Conservatorship had become available. ¹⁶ Goodsir was appointed and left Anstruther. Moving to Edinburgh, he planned to separate the strictly limited teaching component of the University Anatomy collection from the wider material and its use. He began lecturing during the week and giving Saturday morning demonstrations, with an emphasis on pathological anatomy.^17,18

Then in 1841, the position of Conservator to the museum of the RCSEd became vacant when William MacGillivray moved to Aberdeen. ¹⁹ Chosen by the Curators, Goodsir succeeded MacGillivray on 21 April (Figure 1). In addition to caring for the museum collections, the College benefitted from the lectures they had invited him to give. Two years later, Professor Syme offered Goodsir the curatorship of the whole University collection at an annual salary of £150. Goodsir accepted the offer and resigned from the RCSEd. He was followed as College Conservator by Harry who continued until 1845. OPEN IN VIEWER

In May 1844, Goodsir was appointed Anatomy Demonstrator to Professor Monro tertius and made proposals for the future needs of the Medical School Departments. By 1845, he was able to publish Anatomical and Pathological Observations, ²⁰ three chapters by Harry. The contents of the book, summarised by Cameron, ²¹ based on lectures given at the RCSEd in 1842–1843, attracted international recognition. Meanwhile, Goodsir’s presentations to a variety of Societies, the majority published, continued unabated. Many are reproduced in Turner’s book, ¹ together with some not previously in print. He planned but did not finish a System of Dissections and edited an edition of Hannover’s book on the microscope. ²² Later, in 1863, he was invited to assist David Brewster with an article for the North British Review on Faivre’s analysis of Goethe’s studies. ²³ Sir David Brewster, a distinguished scientist, had by that time become Principal of Edinburgh University.

Cell theory

Goodsir continued enthusiastically to study a wide variety of organisms, with the help of the advances in compound microscope design that had taken place ⁴ (Figure 2). The Microscopical Society of London had been formed. ²⁴ Fifteen of his papers describing a variety of organisms were quoted by Lonsdale. ¹ On the basis of this research he concluded that all living organisms are formed of microscopic units, cells. ³ Goodsir was not alone in postulating such a concept and the discovery of cells in plants and animals has been attributed to Schleiden ²⁵ and to Schwann. ²⁶ OPEN IN VIEWER

In the first short chapter of Anatomical and Pathological Observations Goodsir described the microscopic components of the textures, the tissues, as centres of nutrition (Figure 3). They were germinal spots of which one example was that of the ovum, and came to be known as cells. He believed that these circumscribed units with their nuclei drew nutritive substances from nearby capillaries, distributing them to the organs and tissues. Aggregates of centres of nutrition formed germinal centres. Organisms appeared to consist of departments, each comprising simple or developed cells, all related to a single ‘central’ (capital) cell around which the others were grouped. In addition to these components of the tissues and organs were germinal membranes, the parent structures of glands. One surface of each germinal membrane was in contact with a subjacent capillary network, the other giving rise to daughter cells from the most superficial of which came the secreted, glandular products. OPEN IN VIEWER

Goodsir chose the intestinal villi as examples of germinal structures. Within 3 hours of feeding a dog, the lacteals were distended with lymph, the intestine filled with fat globules and desquamated epithelial cells, some derived from the lining of the intestinal villi. The villi were now bare except at their bases where a membrane continued to link their extremities with the intestinal crypts of Lieberkühn. Within the intestinal glands were two varieties of cell: the germinal (‘nutritive’) cells which obtained the substances essential for new cell formation from subjacent capillaries; and the secretory, those needed to synthesise secretions such as bile. The development, maturation and atrophy of the cells of a gland Goodsir found to be proportional to its growth and secretory activity. Cell growth and cell secretion differed only in that in the latter process part of the synthesised matrix was ejected. Whether glandular cells secreted bile, milk or some other product, they were in essence of an identical structure: each was formed of a cell wall, a cavity and a nucleus. Glandular acini originated from single cells each of which had the property of absorbing the particular substances necessary for growth, and subsequently modifying them. In the same manner, each cell species appeared to have a unique survival time, ‘peculiarity of constitution’, ‘irritability’ and ‘morbid action’. These characteristics could be recognised in processes such as ulceration.

Goodsir turned to serous membranes. Considering inflammation, he described how an exudate of serum or lymph passes from subjacent capillaries to the free serous surface, questioning whether these vessels may acquire the capacity for diluting or thinning out exudates during inflammation. Goodsir continued by reviewing the structure of the lymphatic glands into which afferent lymphatic channels enter or from which efferent vessels leave. A lymph gland appeared to be a network of lymphatic vessels supplied by a web of capillaries. A description of the histology of the human placenta followed before Goodsir wrote of bone. The cells that lay between osseous blood vessels and the wall of the Haversian canals originated from the cartilaginous matrix from which the bone took origin. He concluded by analysing the regeneration of bone that follows injury.

These discoveries led him to ask: ‘What is a cell with its walls, contents, nucleus and nucleolus? How is a cell formed? How do cells multiply?’ Evidence supporting his views came, for example, from his 1841 observations on the sunfish. ²⁷ By 1842, his lectures began to contain further ideas on cell theory and his description of Sarcina ventriculi offers one example. ²⁸ Goodsir viewed cells as the structural and functional entities shared by animals and plants. However, he did not claim that cell division and multiplication were attributable to their nuclei nor did his observations provide definitive evidence for the importance of cells in inflammation. Much remained to be discovered by Rudolf Virchow, among the first to acknowledge the importance of Goodsir’s original work. ⁵

Professor of anatomy

When Alexander Monro tertius retired in 1846 from the Edinburgh University Chair of Anatomy, Goodsir applied for the position. An important consideration for the Council was the Presbyterian orthodoxy of applicants. ²⁹ Goodsir fulfilled this and their other demands and was appointed.

Always keen to use his past experience in order to practice as a surgeon and in this way to supplement his income, in 1847 Goodsir moved to the house at 55 George Square, Edinburgh where his brother Joseph lived. Anticipating success, in 1848 he applied for the vacant position of Assistant Surgeon to the Royal Infirmary of Edinburgh. Deeply disappointed when he was not chosen, he ended his relations with the hospital and moved house again before finally settling in Edward Forbes’ South Cottage at Wardie. The cottage was close to the shore of the Firth of Forth and it was here that he lived for the last 10 years of his life.

Goodsir’s working hours extended throughout the day and evening. He added anatomical and pathological material to his own natural history collection and, ambitiously, aimed to rival the unique collections in London of the Hunterian Museum of the Royal College of Surgeons of England, the RCSEng. Goodsir’s purpose was to restore the reputation of the University Department of Anatomy to the admired position it had held in the days of the first two Monros. A quiet man, when disposed to speak or to reply to a question, Goodsir’s knowledge and wisdom were at once apparent, his ideas precise and relevant. His lectures to students were popular but it was inevitable that he was less prolific with his public speaking. The style he assumed may be deduced from a lecture introducing his first course in the winter of 1846. ³⁰ Over the succeeding 5 years, he sought to improve the work of the anatomy dissecting rooms. As Lonsdale wrote, ‘Goodsir did not simply give facts: he illuminated them’. Goodsir (Figure 4) introduced tutorial teaching and for some time assumed personal responsibility for the microscopic demonstrations. By 1860–1861 the size of his class had grown to 354. He encouraged students to compete for an Anatomy gold medal. Winning this medal was undoubtedly a factor enabling James Bell Pettigrew, his best student, to gain the appointment as Assistant to the Museum of the RCSEng. ³¹ OPEN IN VIEWER

In 1846, Goodsir addressed the Royal Society of London On the supra-renal, thymus and thyroid bodies ³² in a paper read by Richard Owen of whom he was a fervent admirer. Elected to Fellowship of the Society, Goodsir continued to contribute to the Royal Society of Edinburgh and to the BA. A potato disease, ‘potato blight’, was wreaking havoc with nutritional demands in the United Kingdom and even more severely in Ireland. Goodsir spoke to the Botanical Society of Edinburgh, the BSEd, on this subject, ³³ viewing the disease as an epidemic and associating its spread with the presence of an organism, Botrytis infestans, which he had detected microscopically. He admired horses and in 1849 presented a cast of a dissected animal to the Royal Institution in Edinburgh. He investigated the changes caused to an elephant tusk by the impact of a musket ball ³⁴ and later, dissected one of these animals. ³⁵

Societies

Goodsir’s interests drew him to many societies and he served as President of every one to which he was elected except the Royal Society of London. In 1833, aged 19, he joined the Edinburgh Royal Medical Society, the RMS. It was the year in which he became a member of the APS but it was not until 1840 that he felt able to take an active part in RMS debates before becoming Senior President for 1841–1842. He was elected as a member of the Literary and Philosophical Society of St Andrews in 1838, the year in which this body was established, but transferred to honorary membership after his move to Edinburgh. He joined Jameson’s Wernerian Society and in 1841, the year in which the RPS welcomed him, the BSEd. In 1848, while chairing the University Veterinary Department and examining Professor Dick’s students sitting the Veterinary diploma, he became a member of the Highland and Agricultural Society. It was only ill health in 1866 that ended his time as an examiner. In 1850 he presided over the sections of physiology and zoology at a meeting of the BA and joined the Aesthetic Club, formed in 1851, promoted, among others, by JY Simpson, by Dr John Brown, renowned for Rab and his Friends, as well as by artists and lawyers.

Neurology, morphology and the geometry of joints

Closely familiar with Goethe’s view on the ‘variety of organic composition’, Goodsir came to believe that morphology complemented chemistry and physics in the study of anatomy. He emphasised that in investigating the human skeleton, the relations of the vascular and nervous systems must be considered, reasoning that the upper limb because of its nerve supply, was an appendage of the lower part of the neck ³⁶ and not, developmentally, a part of the head.

Considering the morphology of limbs, the distribution of the nerve supply could help to understand the way in which different organisms were formed. Among his conclusions was the claim that a limb does not necessarily derive its elements from one somatotome. ³⁷ He compared the location and distribution of the ganglia in the nervous systems of annulose (segmented) organisms with those in vertebrates, ³⁸ studies associated with his analysis of the vertebrate head. ³⁹

Goodsir’s early interest in mechanics led him to a later study of the geometry of living organisms. ⁴⁰ His analysis of skeletal muscle revealed fasciculi in which the common pattern was triangular. In bone, the arrangement of the Haversian canals shared the same form. His favourite speculation was that the triangle was the ‘ground plan’ of all organic forms, bringing all living organisms into the same perspective as crystals, the triangle the mathematical form on which both the organic and the inorganic worlds originated. There was an analogy with objects like the pyramids and buildings of ancient Egypt. In a practical demonstration of how the triangle could explain all living structures, he used the shape and varied positions of his hands and contrasted the morphology of trees with that of the human body. He extended his theory to spindle-shaped cells, each with a ‘parent’ cell, arguing that Man was a conglomerate of cells which matured and decayed like protozoa. He viewed man as a tetrahedron. Following the work of John Leslie ⁴¹ and the study of Nautilus, marine molluscs with a hair-like covering, he suggested that ‘logarithmic spirals’ were the forms governing the increase of organic bodies.

Goodsir developed a particular interest in joints and suggested that the articular surfaces of a bone such as the patella could be viewed as comprising different areas and facets, each performing its individual function during movement.^42,43 In the knee, Goodsir believed that the gliding and rolling movements of flexion and extension were performed between a combination of anterior and posterior, conical, double-headed screws. He contributed a paper on the mechanics of joints ⁴⁴ and proposed that the ‘folds’ of the synovia ‘act as stuffing pads, passing into the spaces left during action while smearing the synovia over the opposing cartilaginous surfaces’. Goodsir concluded that the movements of joints were spiral (helical), an idea, among others, that stimulated James Bell Pettigrew’s later concept of the universality of such movements. ⁴⁵

Goodsir was drawn to the anatomical significance of the units of the nervous system. He exploited his interest in nerve endings, in the relations of nerve fibres to nerve cells and to the special senses. ⁴⁶ In 1839 he had observed the eyes of a mollusc ⁴⁷ and, subsequently, pursued the properties of the retina, ⁴⁸ the manner in which the eye adjusts to distinct, distant vision ⁴⁹ and the contrast between ‘simple’ and ‘compound’ eyes. ⁵⁰

Goodsir’s philosophy

Goodsir believed that Man consisted of corporeal, psychical and spiritual elements, each originating in a northern, temperate region. The Anatomy lectures on The Dignity of the Human Body that he gave in 1862–1863 ⁵¹ represented his views on the Nature of Man and the relative significance of animal life. Goodsir claimed that life was in two forms: Man and All forms below Man. Every species of animal he considered to have been created solely for a geographical area to which it was adapted. Man, Goodsir believed, had not been created for any specific region or climate. Animal mental processes were simple, direct and could not be transgressed. By contrast, the human economy was under the control of his or her own experience so that a human being, but not an animal, could act or not as he or she pleased.

Goodsir then elaborated on the theme that the human body, unlike the ape’s, was a perfect structure in accordance with a ‘higher conscious’ or spiritual principle. Although Man often neglected this principle, making unfortunate or dangerous decisions, these choices, like the occurrence of disease, did not contravene the underlying principle of the perfection of human structure. Whereas the animal body, Goodsir concluded, was complete in its purpose, it was incomplete morphologically. By contrast, the human body was complete in both respects. Man, it was held, had been created separately in God’s image, fully formed and not subject to evolution. The human brain was superior in its geometrical proportions, he believed, to the brain of animals, a superiority comparable to that of the bones, joints and organs of sense. The symmetry of the brain related to the higher faculties. Speech, Goodsir thought, was conferred by an unconscious Divine process over which man had no control. He hoped that his opinions would help to check the spread of Darwinism in England ⁵² and counteract Huxley’s lectures on ‘The relation of man to the lower animals’. ⁵³

Later years

By 1850 Goodsir was unwell. Despite his own ill health, when, in 1853, Professor Jameson became ill, Goodsir adopted the responsibility for his lectures but instead of following his curriculum, presented original addresses of his own. After giving these lectures, Goodsir was exhausted, ‘shrunk in features, worn in body, shattered in nerves and almost a helpless invalid’. He decided on a period of rest and in August 1853 journeyed to Wildblat in the Black Forest in Germany to seek the advice of consultants in Frankfurt-am-Main and Göttingen. After spending the winter in Nice, he returned home in the autumn of 1854 only to suffer the death of his friend Edward Forbes in November. Without hesitation, Goodsir assumed Forbes’s lectures on Natural History and in 1856 was able to address the RMS on the manner in which physiological subjects tend to converge. ⁵⁴

In 1857, Goodsir travelled to Berlin and encountered Johannes Müller and Du Bois. It was the year in which he visited Acland in Oxford, a visit leading him to hope that the Oxford system of education might be emulated in Edinburgh. In 1859 Goodsir acted once more as Promoter at the graduation of Edinburgh medical students. ⁵⁵ He gave the Presidential address to the Medico-Chirurgical Society, ⁵⁶ visited Paris and, in the autumn, Leipzig, Dresden, Vienna and Prague

In 1863 Goodsir welcomed Charles Darwin who had come to consult him about a ‘serious stomach ailment’, a visit presumably prompted by Goodsir’s work on Sarcina ventriculi. ²⁸ In the winter session of 1866, Goodsir started his lecture programme as usual but illness took hold and one day he collapsed in front of his class. A recluse, he now avoided visitors by starting work at 5 o’clock in the morning, ending at 8.30. He fought against disease but became increasingly weak physically and is said to have ‘used his body like a machine’. ⁵⁷ In his lonely cottage, his sister Jane read to him and played the harmonium. She wrote that his last few weeks were spent in bed with the works of Isaac Newton, a Bible and a work on crystallography together with models illustrating his planned publication ‘on the triangular basis of organic forms’.

John Goodsir died at South Cottage on 6 March 1867, aged 52.^58–63 It was the cottage where his friend Forbes had died in 1854. Turner ⁶¹ recorded ‘the pain and infirmity’ of Goodsir’s ‘enfeebled state of health’. Creighton ² wrote of ‘a long illness with wasting of the spinal cord’. Millar ⁶ commented ‘the disease which had attacked him although apparently arrested for a time, made slow and insidious progress and assumed the characters of locomotor ataxia’. An autopsy was performed by Dr Chiene and Mr Stirling. The cause of death, registered on 26 March 1867, begins ‘Atrophy of Spinal Cord’ but the changes described are suggestive of those attributable to a prolonged deficiency of vitamin B₁₂.

After his death, a mass of his notebooks and incomplete manuscripts on anatomy and other subjects was found. At his funeral there were many University professors, Fellows of the RCSEd and friends, together with 200 students. Goodsir was interred in Edinburgh, at the Dean cemetery. One side of his gravestone, engraved symbolically with a spiral, curved line, was beside that of Edward Forbes on which are inscribed the dates 12 February 1815 to 18 November 1854. The University Senatus Academicus adopted a Minute recording the passing of a ‘distinguished man of science with a European reputation’.