Historical Records of Australian Science
, 2013, 24
Mollie Elizabeth Holman 1930–2010
Elspeth M. McLachlan
A,C and G. David S. Hirst
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
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
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 . 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
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  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 . 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 , 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  until
a large range of purine receptors was identified
by Burnstock and others.
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
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 . 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
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 , Ed Daniel , Joe Szurszewski
that put together the first volume of the Pro-
[33, 34] and Gerry Silverberg from the USA,
of the Society, published in 1970, and
Hiroshi Kuriyama , Tadao Tomita  and
was elected to Honorary Membership after her
Y. Hashimoto [23, 26, 27] from Japan, Tom Muir
 and Hugh McKirdy  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 , 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
. 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),
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
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.
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
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.
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
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-
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