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Historical Records of Australian Science, 2013, 24, 242–
Mollie Elizabeth Holman 1930–2010
Elspeth M. McLachlanA,C and G. David S. HirstB A Neuroscience Research Australia, Randwick, NSW 2031, Australia. B86 Caroline Street, South Yarra, Vic. 3141, Australia. CCorresponding author. Ema Mollie Holman was a biophysicist whose work on the autonomic nervous system and the innervation of smooth muscle was seminal in advancing knowledge of its behaviour at a cellular level. She was particularly known for her technical expertise in microelectrode recording of membrane potential from single smooth muscle cells, and the interpretation of their electrical activity, both spontaneous and in response to transmitters released from their autonomic nerves. Mollie Elizabeth Holman was born in Launce- ston, Tasmania, on 18 June 1930 and died in Melbourne on 20 August 2010 after a long illness. Mollie was a distinguished electrophys- iologist who made major contributions to our understanding of the cellular mechanisms under- lying the excitation of smooth muscle and the actions of its innervation by the autonomic ner- vous system. Her work was seminal in the development of knowledge of the functioning of the vascular and visceral tissues of the body. She was involved in training and encouraging a whole generation of autonomic neurobiologists who have led the world in this field for many years. Most of her career was spent at Monash University in Melbourne, which she joined soon after its establishment and where she remained for the rest of her working life. Mollie qualified for a Bachelor of Science in Physics (1951) and Master of Science in Phys- father was a pioneer radiologist who set up the iology (1954) at the University of Melbourne X-Ray Department in the hospital in Launce- and then went to the United Kingdom on a ston. He strongly supported his girls in their Melbourne University Travelling Scholarship to academic pursuits and had a strong influence work on the physiology of smooth muscle at the on Mollie during important times in her life. University of Oxford. She received a DPhil from Starting her primary education in 1938, Mollie Oxford in 1957. Mollie returned to Australia in attended, until the Intermediate year, the girls' 1958 to take up a Lectureship in Physiology at grammar school Broadland House in Launces- the University of Melbourne. In 1963 she moved ton, where she quickly rose to the top of her class. to the Department of Physiology at Monash Uni- She showed the first signs of her future inter- versity. Initially appointed as a Senior Lecturer, ests when, at the age of 14 years, she enrolled she was appointed Reader in 1965 and Professor in evening classes in a course called ‘Intro- in 1970. Mollie retired in 1996. duction to science and engineering'. She loved these studies because ‘it all seemed very logical Early History
and satisfactory'. Mollie moved to Melbourne in Mollie was the eldest of four daughters of 1945 to complete her secondary education as a William Prout Holman and Mollie Bain. Her boarder at Melbourne Church of England Girls' Journal compilation Australian Academy of Science 2013 Mollie Elizabeth Holman 1930–2010 Grammar School (Merton Hall) where she was Oxford to study for a DPhil under the supervi- able to study science to prepare her for matricu- sion of Professor Edith Bülbring in the Depart- lation. She edited the school magazine, played ment of Pharmacology headed by J. H. Burn. basketball and developed a passion for Greek She had been dissuaded by John Eccles (later and Roman history that she kept throughout her a Nobel Laureate) from going to University life. Her chemistry teacher left a great impres- College London where Bernard Katz led the sion, leading Mollie on to study science at the Biophysics Department. Eccles had worked on University of Melbourne. smooth muscle at Oxford with Charles Sher- rington and suggested that this would be a more University Studies
novel area than currently fashionable studies of the skeletal neuromuscular junction. In her first year at the University of Melbourne, Edith Bülbring was a formidable lady and Mollie was based in Mildura at a spillover cam- Mollie and she did not always agree. Bülbring pus for the additional students entering the uni- had had a long, successful career as a classi- versity in the immediate postwar years. While cal pharmacologist using smooth muscle tissues enrolled in Science, she took the subjects for but had developed an interest in smooth muscle the first year of the medical course plus Pure physiology. The field of smooth muscle research Mathematics, which gave her a strong basis for was in its infancy. Since smooth muscles form her later career. She graduated with Honours in the major part of blood vessel walls, dysfunction Physics in 1951, having also studied Physiology leads to disorders of blood pressure and flow. and Biochemistry in her final year, and gained Similarly the intestines, reproductive organs, a Melbourne University scholarship for further bronchioles and urinary tract organs all depend studies. She joined the Department of Physiol- upon the ordered functioning of the smooth mus- ogy and Pharmacology, then under the leadership cles present in their walls. It was also known that of R. D. Wright (a friend of her father, who had by the activity of smooth muscles was controlled by then moved to Melbourne to be head of the radio- the electrical gradient across their cellular wall, therapy unit at the new Peter MacCallum Cancer termed their membrane potential. Hospital) and studied biophysics, a very new At the time of Mollie's arrival in Oxford, subject at the time. Her supervisor was Frank the conventional wisdom was that membrane Shaw (to become the first Professor of Pharma- potentials could only be measured using sharp cology at Melbourne in 1954) who suggested that glass microelectrodes filled with a conducting she measure the voltage generated by sodium solution. These were very fine micro-pipettes: ions pumped across frog skin—subsequently a narrow tubing was heated in a flame until it was classical model for epithelial transport [1]. The white hot and then pulled to make a very fine cap- project involved building her own electrodes and illary tip that was hopefully patent. The pipette recording apparatus. In 1953 and 1954, Holman was filled with concentrated potassium chloride was a demonstrator in Pharmacology as well as a conducting solution—if you were lucky and as working on her research. She completed an the tip of the electrodes did not break off when MSc at the University of Melbourne in Septem- the pipettes were boiled under reduced pres- ber1954 with a thesis entitled ‘Pharmacology of sure in concentrated potassium chloride solution. Bioelectricity'. She had given her first scientific The sharpness of the electrode was defined paper at an ANZAAS meeting in Canberra at the by its tip diameter which, since it was too beginning of 1954. small to be viewed in the light microscope, During this time, she started to read about was assessed by measuring its resistance—the the autonomic nervous system. This system con- finer the tip, the higher the resistance. At the sists of the nerves that control all the organs time, the conventional wisdom was that any and tissues of the body except skeletal muscles, electrodes with resistances above 10 MQ were a subject that fascinated her for the rest of her life. likely to be incompletely filled and therefore Pushing such electrodes into smooth mus- cle resulted in resting (trans)membrane poten- Since it was unusual to carry out postgraduate tials of about −10 mV (inside relative to the studies in Australia at that time, Mollie went to Historical Records of Australian Science, Volume 24 Number 2 external solution), with action potentials having achievements but she was immensely respected amplitudes of some 5 to 7 mV. Action poten- by physiologists and clinicians alike. tials are brief positive-going potentials exhibited While in Oxford, Mollie befriended Mike by all excitable cells—such as nerve and mus- Rand, an Australian who was working with cle cells. Edith Bülbring had published several J. H. Burn and was soon to become the sec- papers showing such records. Moreover, as the ond Professor of Pharmacology at the University discharge of action potentials was suppressed by of Melbourne) and Geoffrey Burnstock (a post- the removal of Na+ ions, it was thought that doctoral fellow in Edith Bülbring's laboratory). smooth-muscle action potentials resulted from Geoff was originally from London and was soon an inward movement of Na+ ions. Mollie rea- to become Professor of Zoology at the University soned that given the small diameters of smooth muscle cells (2–3 μm diameter), the electrodes being used were likely to damage the cells Return to Melbourne
and give unreliable measurements. She therefore developed a protocol for manufacturing elec- After graduating at Oxford in 1957, Mollie trodes with much finer tips; these had resistances was offered a lectureship in R. D. Wright's of some 60 to 80 MQ. With these, she found Department of Physiology and went back to that the resting membrane potentials of smooth- the University of Melbourne where she set up muscle cells were, as in many other excitable her own laboratory in the departmental paint cells, around −70 mV [3–5]. Further, the peak of store. She started experiments, initially using the action potential often overshot 0 mV (that is, equipment borrowed from David Dewhurst until their amplitude was nearly 100 mV), and chang- she obtained grant funding for newer record- ing the ionic concentrations (as she had done ing apparatus. She developed her collaboration with frog skin) indicated that the ions entering with Geoff Burnstock to work on the electro- the cells were Ca2+, not Na+. physiology of neuromuscular transmission in the At the time, these findings were greeted with guinea-pig vas deferens. At that time it was general disbelief: Mollie had overturned many widely held that the autonomic nervous system of the recent findings made in the Oxford labo- lacked sophistication: the sympathetic nervous ratory. Furthermore she had used electrodes that system was thought to be switched on in much were thought to be unreliable. The day was car- the same way as one would turn on a tap. By ried, however, by two of Mollie's most charming recording the membrane potential of the smooth attributes. First, she was modest and encour- muscle cells during electrical stimulation of aged discussion about her possible failings. Sec- the sympathetic postganglionic nerves, Mollie ond, rather than hoarding her new technical identified excitatory junction potentials (ejp's) skills, she shared them with others, in particular that she showed were analagous to the endplate her Japanese colleagues Hiroshi Kuriyama and potentials recorded at the neuromuscular junc- Tadao Tomita, who rapidly validated her obser- tion and excitatory postsynaptic potentials in the vations. These aspects of Mollie's character led central nervous system [6, 7]. Mollie's studies on both to rapid growth in understanding of smooth the vas deferens revealed that the nerves released muscle electrophysiology and the generation of quanta of transmitter [9] and the effects of these a fine spirit of camaraderie amongst smooth- summed to give rise to post-junctional excita- muscle workers from all continents that persists tory potentials in the smooth muscle cells. If sufficient synaptic activity occurred, the poten- Mollie's success in correcting Bulbring's tials summed to trigger action potentials, whilst observations on membrane potentials and action low-level activity demonstrated patterns of facil- potentials in smooth muscle was the talk of itation detected elsewhere [16]. Once again the Oxford biological sciences. It also gained her conventional thinking had been overturned and considerable esteem within Britain's Physiolog- great progress made. Mollie and Geoff Burn- ical Society. She later became an international stock published a series of pioneering papers leader who was highly regarded in Europe and describing this work [6–11, 13, 14, 16, 17]. North America as well as in Britain, Aus- A major puzzle was that the responses to sym- tralia and Japan. She did not broadcast her pathetic nerve stimulation were not blocked by Mollie Elizabeth Holman 1930–2010 adrenoceptor antagonists [17], which was sur- prising since noradrenaline was well established as the neurotransmitter released by sympathetic nerves, and adrenoceptor antagonists block most of the contractile responses. It was not until over twenty years later that Burnstock and several other groups showed that the ejp's were the mem- brane's potential change in response to adenosine trisphosphate (ATP) released as a co-transmitter with noradrenaline from sympathetic nerves. During this time, Mollie and Geoff collab- orated with Graeme Campbell and Max Ben- nett, then postgraduate students of Burnstock, to discover the non-adrenergic, non-cholinergic (NANC) inhibitory innervation of the guinea-pig taenia coli [12, 15]. Later Mollie came to accept that ATP is involved as a co-transmitter in this and other autonomic systems, although she (and many others) resisted this idea for years [51] until a large range of purine receptors was identified by Burnstock and others. Figure 1. Mollie Holman at the dissecting micro-
scope preparing smooth muscle tissue for electro- physiological recording in vitro, in the Department Research at Monash
of Physiology at Monash University in 1978 (image by Herve Alleaume, courtesy of Monash University In 1963 Mollie moved to the Department of Archives IN268). Physiology at Monash University, enticed by Archie McIntyre, FAA, the foundation profes- sor and a major driving force behind Australia's to the colon that did not involve the spinal cord. excellence in neurophysiological research. Ini- Other studies led to an understanding of the prop- tially appointed as a Senior Lecturer, Mollie erties of myenteric neurons in the enteric ganglia was appointed Reader in 1965 and to a personal and, for example, the prolonged afterhyperpolar- chair in Physiology in 1970. Much of Monash's izations that limit the firing properties of these international reputation in biological and med- and many other types of neuron, including in the ical science in the 1970s and 1980s was due central nervous system [38, 39] (Fig. 1). to her scientific accomplishments. In 1965, she At Monash in the 1970s and into the 1980s, was awarded the Edgeworth David Medal by Mollie established and led the Neuropharma- the Royal Society of New South Wales for her cology Group, which included Robert Bywater, achievements as a scientist aged under 35, and David Hirst, Elspeth McLachlan and Grahame in 1970 Monash awarded her a Doctor of Science Taylor. The Group was rather special, being com- posed of academics who were all more or less Mollie collaborated with Gary Blackman at interested in the same things, namely smooth the University of Otago in New Zealand to use muscle and other targets of the autonomic ner- microelectrodes to study the synapses in auto- vous system and the nerve pathways themselves. nomic ganglia [31]. They extended and improved It was a rather unusual situation to have so many the first recordings made by others by using individual academics with overlapping interests finer-tipped electrodes and applied these in intact and with the ability to be independent at the same preparations of ganglia and smooth muscles. At time as being part of the group. It was not at all her initiation, studies were begun on ganglia like the modern groups working under a senior exerting control over gastrointestinal motility scientist. Mollie gave everyone encouragement [34, 35]. These studies led to the first unequivo- to work on what they wanted and she supported cal demonstration of a peripheral reflex between us all. We met weekly to discuss our results intestinal afferents and sympathetic projections and problems that we needed help to solve. Historical Records of Australian Science, Volume 24 Number 2 Often, someone would come up with a com- the most exciting and productive department of pletely different approach—sometimes a techni- physiology in Australia at that time. cal idea, sometimes an intellectual solution. And Mollie was present as a group leader, much like a cheerleader providing drive and inspiration but Outside the Laboratory
not directing, to help us achieve our goals. She Mollie was active as a member of the Australian pointed us to relevant literature and passed on Physiological Society, presenting many papers the latest developments she had heard about at at its meetings and serving terms as National meetings overseas. Secretary and later as President (when it was There were many notable international the Australian Physiological and Pharmacolog- visitors to the Group at that period: Ladd ical Society). She led the editorial committee Prosser [11], Ed Daniel [43], Joe Szurszewski that put together the first volume of the Pro- [33, 34] and Gerry Silverberg from the USA, ceedings of the Society, published in 1970, and Hiroshi Kuriyama [16], Tadao Tomita [41] and was elected to Honorary Membership after her Y. Hashimoto [23, 26, 27] from Japan, Tom Muir [33] and Hugh McKirdy [39] from Glasgow, Mollie was also involved in significant Donald Cheung from Canada, Nick Spitzer and roles within the Australian and New Zealand Alan North from the USA, and Wilfrid Jänig Association for the Advancement of Science from Germany. There were also many students (ANZAAS), served on the Victorian Board of the and postdoctoral fellows, including Annmarie National Heart Foundation (1975–7) and took Surprenant [45, 47, 48, 50, 52], Tim Neild on major responsibilities with research organiza- [46, 50, 52], Rick Lang [53], Helena Parking- tions like the Australian Research Grants Com- ton, Harry Coleman and Mary Tonta [55–58], mittee, forerunner of today'sAustralian Research Terry Smith, Susan Luff and Marianne Tare, Council. She was the first female member of some of whom later joined the academic staff the Executive of the Commonwealth Scientific at Monash or other universities internationally. and Industrial Organisation (1975–7). In all these A symposium held in Mollie's honour in 1992 at activities she was never the token female—she Great Keppel Island, Queensland, was attended fully embraced her responsibilities and worked by most of her Australian and international tirelessly in the promotion of the highest levels of scientific integrity. From the late 1980s, Mollie was less involved In 1985, Mollie was awarded the ANZAAS in the laboratory although she continued to play Medal, for services in the advancement of sci- the same supportive role for the members of ence or administration and organization of sci- the Group. In the 1990s she did some interest- entific activities, or the teaching of science ing experiments with Parkington and Coleman, throughout Australia and New Zealand and in recording from cells in the adrenal medulla that contributions to science that lie beyond normal had many of the properties of neurons in sym- professional activities. The medal was presented pathetic ganglia [55, 56, 58]. Later, her interest by Sir Edmund Hillary (Fig. 2). in the control of the gastrointestinal tract by the Mollie was elected a Fellow of the Australian enteric ganglia led to her involvement in studies Academy of Science in 1970 and served on of the inhibitory regulation of the mouse colon several of the Academy's committees including with Nick Spencer, a student with Bywater and the Biophysics and Pharmacology Group (1968– Taylor [59, 60]. A dinner attended by many of 82, Chair 1973 and 1979), and what was then Mollie's past colleagues was held in Melbourne numbered Sectional Committee 6, the sectional in 2007 to celebrate the fiftieth anniversary of committee covering her field of science (1972– her first publication in the Journal of Physiol- 6, including Chair 1974–5; 1984–7, including ogy. The dinner was organized by Alan North, Chair 1985–7). She also served on the Academy's by then at the University of Manchester and Council (1980–3) and was Vice-President (Bio- President of the Physiological Society. logical Sciences) in 1982–3. During Mollie's years at Monash, Archie Within the Physiology Department at Monash, McIntyre, FAA, and then Bob Porter FAA, were Mollie's involvement in teaching was focused for Heads of Physiology and they and Mollie led many years on running the Honours course, for


Mollie Elizabeth Holman 1930–2010 Figure 2. Sir Edmund Hillary presenting Mollie Holman with the ANZAAS Medal in 1985 (cropped from
original image, courtesy of Monash University Archives IN1321).
which she used her experiences at Oxford as a model. She was a strong mentor to all Physi- ology graduate students over the course of her career. She later spent a great deal of her time in research administration, including a period as Associate Dean of Research of the Faculty of Medicine. She retired from Monash in 1996. Mollie was awarded an Honorary Doctor of Laws degree by Monash in 1999. In 2007 she received the David de Kretser Award for her exceptional contribution to the University (Fig. 3). Mollie's great strength, aside from her excel- lence as a scientist, was her ability to support and encourage others. With each visitor, Mollie provided amazing support. With each new staff member, whatever their field of study, Mollie ensured that their research efforts could proceed Figure 3. Mollie Holman in 2007 on the occasion
without interference. Most notably, she sup- when she received the David de Kretser Award for ported young scientists. She would seek them out her exceptional contribution to Monash University and aid them in their studies. At scientific meet- (image from Monash University News). ings, both in Australia and overseas, she would invariably be seen encouraging younger work- awarded each year to the best PhD thesis in each ers, promoting their efforts. Her contributions in this way were recognized by Monash by Mollie was appointed an Officer of the Order the creation of the Mollie Holman Medals, of Australia (AO) in 1998. Historical Records of Australian Science, Volume 24 Number 2 Life beyond Science
the guinea-pig's taenia coli. J. Physiol. 136 (1957),
569–584.
Mollie had many interests outside her science 4. Holman, M. E. Membrane potentials recorded and the academic life. She was a hiker, an with high-resistance micro-electrodes; and the accomplished skier and an intrepid traveller, par- effects of changes in ionic environment on the ticularly in India and the Middle East, with many electrical and mechanical activity of the smooth stories to tell on her return to Melbourne. She muscle of the taenia coli of the guinea-pig. J. Phys- also loved Greek and Roman history and did iol. 141 (1958), 464–488.
trips to several archaeological sites. As well, she 5. Bülbring, E., Burnstock, G., and Holman, M. E. enjoyed painting in watercolour. Excitation and conduction in the smooth mus- Mollie suffered a couple of strokes in the cle of the isolated taenia coli of the guinea-pig. 1990s from which she determinedly recovered. J. Physiol. 142 (1958), 420–437.
6. Burnstock, G., and Holman, M. E. Autonomic After one of these, she started driving again after nerve-smooth muscle transmission. Nature 187
the minimum three months, much to the dismay (1960), 951–952. of some of us as she was only just beginning to 7. Burnstock, G., and Holman, M. E. The walk without a stick. She continued her inter- transmis- sion of excitation from autonomic nerve ests in travel and history and was always keen to smooth muscle. J. Physiol. 155 (1961), 115–
to hear the scientific gossip when any of her old colleagues visited her. 8. Burnstock, G., and Holman, M. E. Effect of dener- Just a few weeks before Mollie died, she was vation and of reserpine treatment on transmission mentally as bright and interested as ever and at sympathetic nerve endings. J. Physiol.
160 (1962), 461–469.
insisted on winning her hand of Scrabble before 9. Burnstock, G., and Holman, M. E. Spontaneous she would chat with visitors. It was a shock to potentials at sympathetic nerve endings in smooth hear that she died so soon afterwards. Mollie was muscle. J. Physiol. 160 (1962), 446–460.
a peerless scientist and her lasting memory is 10. Burnstock, G., and Holman, M. E. Smooth mus- one of generosity of spirit, her support for the cle: autonomic nerve transmission. Annu. Rev. young and her sense of a fair go for all. She Physiol. 25 (1963), 61–90.
leaves a great scientific legacy and many friends 11. Burnstock, G., Holman, M. E., and Prosser, and colleagues who still treasure her input to C. L. Electrophysiology of smooth muscle. Phys- iol. Rev. 43 (1963), 482–527.
12. Burnstock, G., Campbell, G., Bennett, M., and Holman, M. E. Inhibition of the smooth muscle
of the taenia coli. Nature 200 (1963), 581–582.
We are very grateful to Robert Porter and Rick 13. Merrillees, N. C., Burnstock, G., and Holman, Lang for their input to this memoir. Details were M. E. Correlation of fine structure and physiol- provided by the CSIRO and the Heart Foundation ogy of the innervation of smooth muscle in the and images by the Monash University Archives. guinea pig vas deferens. J. Cell. Biol. 19 (1963),
The obituary by Professor Uwe Proske published in The Age (13 October 2010) provides further 14. Holman, M. E., and Jowett, A. Some actions of catecholamines on the smooth muscle of the details of Mollie's life and contributions. The guinea-pig vas deferens. Aust. J. Exp. Biol. Med. portrait photograph was taken in the late 1970s Sci. 42 (1964), 40–53.
15. Burnstock, G., Campbell, G., Bennett, M., and Holman, M. E. Innervation of the guinea-pig taenia coli: are there intrinsic inhibitory nerves 1. Holman, M. E., and Shaw, F. H. The effect of which are distinct from sympathetic nerves? Int. yohimbine and other drugs on the isolated J. Neuropharmacol. 3 (1964), 163–166.
frog skin potential. Aust. J. Exp. Biol. Med. 16. Burnstock, G., Holman, M. E., and Kuriyama, Sci. 33 (1955), 671–676.
H. Facilitation of transmission from autonomic 2. Holman, M. E., Johnstone, B. M., Shaw, F. H., and nerve to smooth muscle of guinea-pig vas defer- Simon, S. E. The effect of changes of environ- ens. J. Physiol. 172 (1964), 31–49.
ment on the electrical and ionic pattern of muscle. 17. Burnstock, G., and Holman, M. E. An elec- J. Gen. Physiol. 40 (1956), 263–288.
trophysiological investigation of the actions of 3. Holman, M. E. The effect of changes in sodium some autonomic blocking drugs on transmission chloride concentration on the smooth muscle of in the guinea-pig vas deferens. Br. J. Pharmacol.
Chemother.
23 (1964), 600–612.
Mollie Elizabeth Holman 1930–2010 18. Holman, M. E., and Hughes, J. R. Inhibition of to guinea-pig pelvic ganglia. Br. J. Pharmacol. 41
intestinal smooth muscle. Aust. J. Exp. Biol. Med. (1971), 26–40. Sci. 43 (1965), 277–290.
34. Crowcroft, P. J., Holman, M. E., and Szurszewski, 19. Holman, M. E., and Hughes, J. R. An inhibitory J. H. Excitatory input from the distal colon to component of the response to distension of rat the inferior mesenteric ganglion in the guinea-pig. ileum. Nature 207 (1965), 641–642.
J. Physiol. 219 (1971), 443–461.
20. Burnstock, G., and Holman, M. E. Effect of 35. Holman, M. E., Hirst, G. D., and Spence, I. Prelim- drugs on smooth muscle. Annu. Rev. Pharmacol. inary studies of the neurones of Auerbach's plexus 6 (1966), 129–156.
using intracellular microelectrodes. Aust. J. Exp. 21. Bennett, M. R., Burnstock, G., and Holman, M. E. Biol. Med. Sci. 50 (1972), 795–801.
Transmission from perivascular inhibitory nerves 36. Holman, M. E., and Spitzer, N. C. Action of to the smooth muscle of the guinea-pig taenia coli. botulinum toxin on transmission from sympathetic J. Physiol. 182 (1966), 527–540.
nerves to the vas deferens. Br. J. Pharmacol. 47
22. Burnstock, G., and Holman, M. E. Junction (1973), 431–343. poten- tials at adrenergic synapses. Pharmacol. 37. Holman, M. E. A discussion on recent develop- Rev. 18 (1966), 481–493.
ments in vertebrate smooth muscle physiology. 23. Hashimoto,Y., Holman, M. E., and Tille, J. Electri- A survey of new findings and the discussions aris- cal properties of the smooth muscle membrane of ing during sessions I. II and 3. Philos. Trans. R the guinea-pig vas deferens. J. Physiol. 186 (1966)
Soc. Lond. B Biol. Sci. 265 (1973), 157–165.
38. Hirst, G. D., Holman, M. E., and Spence, I. Two 24. McIntyre, A. K., Holman, M. E., and Veale, types of neurones in the myenteric plexus of duo- J. L. Cortical responses to impulses from single denum in the guinea-pig. J. Physiol. 236 (1974),
Pacinian corpuscles in the cat's hind limb. Exp. Brain Res. 4 (1967), 243–255.
39. Hirst, G. D., Holman, M. E., and McKirdy, H. C. 25. Holman, M. E., and McLean, A. The innervation Two descending nerve pathways activated by dis- of sheep mesenteric veins. J. Physiol. 190
tension of guinea-pig small intestine. J. Physiol. (1967), 55–69. 244 (1975), 113–127.
26. Hashimoto, Y., Holman, M. E., and McLean, A. J. 40. Holman, M. E., and Weinrich, J. P. The effects of Effect of tetrodotoxin on the electrical activity of calcium and magnesium on inhibitory junctional the smooth muscle of the vas deferens. Nature 215
transmission in smooth muscle of guinea pig small (1967), 430–432. intestine. Pflugers Arch. 360 (1975), 109–119.
27. Hashimoto, Y., and Holman, M. E. Effect of man- 41. Holman, M. E., Taylor, G. S., and Tomita, T. Some ganese ions on the electrical activity of mouse vas properties of the smooth muscle of mouse vas deferens. Aust. J. Exp. Biol. Med. Sci. 45 (1967),
deferens. J. Physiol. 266 (1977), 751–764.
42. Holman, M. E., and Hirst, G. D. (1977) Junctional 28. Holman, M. E., Kasby, C. B., Suthers, M. B., and transmission in smooth muscle and the autonomic Wilson, J. A. Some properties of the smooth mus- nervous system. In: E. R. Kandel (ed.), Hand- cle of rabbit portal vein. J. Physiol. 196 (1968),
book of Physiology. Section 1: The Nervous Sys- tem (American Physiological Society, Bethesda), 29. Chapman, J. B., and Holman, M. E. Potassium con- tractures in the ureter of the guinea-pig. Aust. J. 43. Daniel, E. E., Taylor, G. S., Daniel, V. P., and Exp. Biol. Med. Sci. 46 (1968), 503–514.
Holman, M. E. Can nonadrenergic inhibitory vari- 30. Holman, M. E. Electrophysiology of vascu- lar cosities be identified structurally? Can. J. Physiol. smooth muscle. Ergeb. Physiol. 61 (1969),
Pharmacol. 55 (1977), 243–250.
44. Hirst, G. D., and Holman, M. E. Functions of 31. Blackman, J. G., Crowcroft, P. J., Devine, C. E., enteric nerve cells in relation to peristalsis. Nippon Holman, M. E., and Yonemura, K. Heikatsukin Gakkai Zasshi. 14 (1978), 189–200.
Transmis- sion from preganglionic fibres in the 45. Holman, M. E., and Surprenant, A. M. Some hypogastric nerve to peripheral ganglia of male properties of the excitatory junction potentials guinea-pigs. J. Physiol. 201 (1969), 723–743.
recorded from saphenous arteries of rabbits. J. 32. Holman, M. E. Interpretation of the electromyo- Physiol. 287 (1979), 337–351.
gram of the small intestine as recorded with exter- 46. Holman, M. E., and Neild, T. O. Membrane prop- nal electrodes. Nihon Heikatsukin Gakkai Zasshi. erties. Br. Med. Bull. 35 (1979), 235–241.
6 (1970), 73–74.
47. Holman, M. E., and Surprenant, A. M. An 33. Holman, M. E., Muir, T. C., Szurszewski, J. H., electrophysiological analysis of the effects of and Yonemura, K. Effect of iontophoretic applica- noradrenaline and alpha-receptor antagonists on tion of cholinergic agonists and their antagonists neuromuscular transmission in mammalian Historical Records of Australian Science, Volume 24 Number 2 muscular arteries. Br. J. Pharmacol. 71 (1980),
splanchnic nerves to the adrenal medulla of guinea-pigs. J. Physiol. 478 (1994), 115–124.
48. Holman, M. E., and Surprenant, A. Effects of 56. Holman, M. E., Tonta, M. A., Coleman, H. A., and tetraethylammonium chloride on sympathetic Parkington, H. C. Currents caused by the spon- neuromuscular transmission in saphenous artery taneous release of quanta of acetylcholine onto of young rabbits. J. Physiol. 305 (1980), 451–465.
chromaffin cells in guinea-pig adrenal medulla. 49. Bywater, R. A., Holman, M. E., and Taylor, Neurosci. Lett. 184 (1995), 75–78.
G. S. Atropine-resistant depolarization in the 57. Holman, M. E., Tonta, M. A., Parkington, H. C., guinea-pig small intestine. J. Physiol. 316 (1981),
and Coleman, H. A. Tetrodotoxin-sensitive action potentials in smooth muscle of mouse vas defer- 50. Surprenant, A., Neild, T. O., and Holman, M. E. ens. J. Auton. Nerv. Syst. 52 (1995), 237–240.
Effects of nifedipine on nerve-evoked action 58. Holman, M. E., Tonta, M. A., Coleman, H. A., potentials and consequent contractions in rat tail and Parkington, H. C. Muscarinic receptor artery. Pflugers Arch. 396 (1983), 342–349.
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