93^
Journal and Proceedings
of the
Royal Society
of
New South Wales
2016
Volume 149 Parts 1 &2
Numbers 459 to 462
for the encouragetnent of studies and invest^flons in Science Art Litetalure and Philosophy
The Royal Society of New South Wales
Office Bearers for 2016
His Excellency General The Honourable David Hurley AC DSC (Ret’d)
Governor of New South Wales
Em. Prof. David Brynn Hibbert BSc PhD CChem FRSC FRACI FRSN
Mr John Hardie BSc (Syd), FGS, MACE FRSN
Dr Donald Hector BE(Chem) PhD (Syd) FIChemE FIEAust FAICD FRSN
Ms Judith Wheeldon AM, BS (Wis) MEd (Syd) FACE FRSN
Em. Prof. Robert Marks, BE, MEngSci, ResCert, MS, PhD (Stan.) FRSN
Dr Herma Buttner PhD
Mr Richard Wilmott
Dr Ragbir Bhathal PhD FSAAS
Dr Erik W. Aslaksen MSc (ETH) PhD FRSN
Dr Mohammad Choucair PhD
Prof. Max Crossley PhD FAA FRAC FRSN
Dr Desmond Griffin PhD AM FRSN
Prof. Stephen HiU PhD AM FTSE FRSN
Em. Prof. Heinrich Hora DipPhys Dr.rer.nat DSc FAIP FInstP CPhys
FRSN
Prof. E James Kehoe PhD FRSN
Em. Prof Roy MacLeod AB (Harv) PhD, LittD (Cantab) FSA FALIA
FASSA FRHistS FRSN
Prof. Bmce Milthorpe PhD FRSN
Prof. Ian Sloan AO PhD FAA FRSN
Hon. Prof. Ian Wilkinson FRSN
A/Prof. Chris Bertram PhD FRSN (by invitation)
Mr Hubert Regtop
Executive Office The Association Specialists
Editorial Board
Em. Prof. Robert Marks, BE, MEngSci, ResCert, MS, PhD (Stan.) FRSN - Hon. Editor
Prof. Richard Banati MD PhD FRSN
Prof. Michael Burton BA MA MMaths (Cantab) PhD (Edinb) FASA FAIP FRSN
Dr Donald Hector BE(Chem) PhD (Syd) FIChemE FIEAust FAICD PRSN
Em. Prof. David Brynn Hibbert BSc PhD (Lond) CChem FRSC FRACI FRSN
Dr Michael Lake BSc (Syd) PhD (Syd)
Dr Nick Lomb BSc (Syd) PhD (Syd) FASA FRSA
Prof. Timothy Schmidt BSc (Syd) PhD (Cantab) FRSN
Website: http: / / www.royalsoc.org.au
The Society traces its origin to the ^Philosophical Society of Australasia founded in Sydney in 1821. The Society exists for ‘VA encouragement of
studies and investigations in Science Art Uterature and Philosophy publishing results of scientific investigations in its Journal and Proceedings;
conducting monthly meetings; awarding prizes and medals; and by liaising with other learned societies within Australia and
internationally. Membership is open to any person whose application is acceptable to the Society. Subscriptions for the Journal are also
accepted. The Society welcomes, from members and non-members, manuscripts of research and review articles in aU branches of
science, art, literature and philosophy for publication in the Journal and Proceedings.
Patron
President
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Hon. Secretary (Ed.)
Hon. Secretary (Gen.)
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Councillors
Web Master
Southern Highlands
Branch Representative
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 1-4. ISSN 0035-9173/16/010001-04
Editorial
Robert E. Marks
Finally, the first issue of the JProcRSNSW
under the new editorship. Several
papers: the outgoing presidents address by
Don Hector, the W. B. Clarke Memorial
Lecture by Griffin et ah, the Liversidge Lec¬
ture by Banwell et ah, a paper by the late
Commonwealth Statistician, Ian Castle, on
one of the most illustrious members of the
Royal Society and its forerunner, the Philo¬
sophical Society of N.S.W., the forgotten
polymath, William Stanley Jevons, scientist
and economist, followed by a paper of mine
which puts Jevons’ activities in the Society
into context and updates Castle’s article with
some recent debates about the implications
of Jevons’ work for energy policy. The last
paper is by Foster, a Royal Society Scholar¬
ship winner There are also 16 abstracts of
recent doctoral dissertations from several
N.S.W universities.
The delay in publishing this issue is due to
two things. First, the handover took place
in May, with few accepted papers. Second,
I threw myself into indexing the contents of
past issues back to 1 866 and before, making
the index accessible to Google’s indexing
robots (which has now happened), and
making the contents of past articles readily
accessible to anyone on the Internet.
This process would not have been feasible
without the work done by past editors and
librarians of the Society, the Biodiversity
Heritage Library (the host on the on-line
repository of the Journal), the Smithsonian
Libraries, the Missouri Botanical Garden
Peter H. Raven Library, the Harvard Uni¬
versity Museum of Comparative Zool¬
ogy Ernst Mayr Library, the University of
California Libraries, the Leland Stanford Jr.
University Library, the Wayback Machine’s
Internet Archive, and the National Library
of Australia’s incomparable, but threatened,
Trove on-line archive of Australian mate¬
rial back to first European settlement. For
the on-line repositories of the Journal, past
issues were scanned to PDF and placed on¬
line in large files, one per issue or volume.
These were embedded in a viewing platform
which includes Optical Character Recogni¬
tion output.
Readers can see the format of the main
repository by going to the Journal Archive
page of the Journal’s contents, at the Society’s
web pages.
Clicking on a volume and then a paper
reveals a PDF of each page with a plain¬
text OCR on the right (when clicked on).
Although the files are large, each page (cor¬
responding to a page published in the hard¬
copy version of the Journal) has a unique
URL, which allows us to link the initial page
of the 3,1 10 articles in the Journal smct 1867.
I used plaintext versions of each issue’s con¬
tents to derive the pages of the contents of
each volume. These contents pages, roughly
one a year, enable an on-line index for the
Journal’s contents, with the URLs providing
the links to each paper.
The index of articles and papers starts in
1822, forty years before the Royal Society
was granted its letters patent, when the first
forerunner of the Society, the Philosophical
Society of Australia, was active, under the
patronage of the Governor, Sir Thomas Bris¬
bane. At that time there was no dedicated
publication for the Society’s papers (that
1
Journal & Proceedings of the Royal Society of New South Wales
Marks — Editorial
would have to wait for over thirty years), but
some papers presented to the Society were
published as chapters in a book published in
London in 1825, edited by a former member
of the Society, Barron Field, a lawyer. This
book is now in the public domain and its
contents have been made available on-line
as part of the Gutenberg Project. i
After thirty years of little if any activity,
the Philosophical Society of New South
Wales began in 1856, under the patronage
of the new governor. Sir William Denison,
an engineer. Some papers were published
in the Sydney newspapers, especially Henry
Parkes’ The Empire, and are now available
on Trove. There were two other outlets for
papers in the 1850s and early 1860s: a com¬
mercial monthly. The Sydney Magazine of
Science and Art, published papers from the
Society and other learned groups for the
two years it was in existence. It too is freely
available on-line. The Philosophical Society
of N.S.W. also published the Transactions of
the Philosophical Society ^Tom 1862 to 1865,
and this too is available on-line. In 1867
the Journal of the Royal Society first appeared,
and has done so ever since. Perhaps some¬
one will use this newly accessible resource
to write a paper analysing how the contents,
authorship (numbers, sexes), etc of papers
have changed over the past 110+ years.
In 1955, the president of the Society,
Ronald Nyholm, mused about the three
phases of the Society, as reflected in sub¬
missions to the Journal:
1 The Gutenberg Project s copy appears to come from
the Stanford library, although Stanford University
was only founded in the 1 890s. It turns out at that
Thomas Welton Stanford, brother of Leland Stanford,
the railroad baron who founded Stanford, lived in
Australia for many years and amassed a library of Aus-
traliana, which he bequeathed to the University. (I
thank Jessica Milner Davis for this sleuthing.)
“Broadly speaking, the history of the Soci¬
ety falls into three periods. Before the first
world war the Royal Society of New South
Wales was the main scientific society in
Sydney, at least so far as the physical sci¬
ences were concerned. The Society was, for
scientific people, an important means of
mutual contact, discussion and the Journal
received many of their original researches.
Between the two world wars there were
founded in Sydney many specialist scien¬
tific bodies or branches of older ones, such
as the Institute of Physics and the Royal
Australian Chemical Institute, the meet¬
ings of which catered for discussions of
specialist subjects. Nevertheless, many of
the original papers of these specialist still
found their way to our Journal. Thus, the
1 940 Journal was one of the largest ever,
and in it were 38 research papers. There
were 23 chemistry papers, 4 in mathemat¬
ics and 9 in geology. After the Second
World War we enter the third phase — the
development in Australia of new specialist
journals to cater for the needs of scientists.
Examples of these are the Australian Jour¬
nal of Chemistry and the Australian Journal
of Physics. Furthermore, overseas socie¬
ties publishing specialist journals, e.g. the
Chemical Society of London, speeded up
the rate of handling of papers and went out
of their way to provide air-mail facilities
in order to assist folk submitting papers
from Australia.”
The decline in the number of research papers
received (as distinct from the Presidential
Address, Clarke and Liversidge Lectures) is
shown in Figure 2 of Nyholm’s paper. In
1952 the Journal was the smallest since
1929.
Nyholm argued that this had at least two
effects: the mix of disciplines in the Journal
had become unbalanced, which would even-
2
Journal & Proceedings of the Royal Society of New South Wales
Marks — Editorial
tually affect journal exchanges with other
institutions; and Society members would
lose interest in the Journal and perhaps in
the Society. Sixty years ago, Nyholm quoted
a previous president, Richard Bosworth, who
argued for a policy of encouraging research¬
ers to write reports on their research worded
so as to be intelligible to a novice in the field,
rather than to the expert readers the special¬
ist journals assume.
What was true sixty years ago still holds
today, although the pressure to publish in
A-rated journals means that fewer such
accessible papers will be submitted. And
the fourth phase of the Journal— age
of the Internet— -means that the Journal, at
least in its hard-copy form, is increasingly
an anachronism. The other side of that
coin is the ability to make over 1 60 years of
articles accessible to anyone with a browser.
The Journal still publishes the Clarke and
Liversidge lectures and now also a garland
of papers from the annual forum. We also
publish short abstracts from recent Ph.D.
dissertations. But the flow of first-rate, cut-
ting-edge research papers has stopped long
since.
What is happening at our sister societies?
The Transactions of the Royal Society of South
Australia is the result of an amalgamation
(in 2004) with the Records of the South Aus-
tralian Museum and is published by Taylor
& Francis in both hard and soft copies. Its
2015 impact factor was 0.484. Contents
of the Transactions are not freely available.
Subscriptions to the two annual issues of the
Transactions are included in members’ dues,
but non-members and institutions pay up to
$285 a year for hard and soft copies.
The Proceedings of the Royal Society of Victo¬
ria is published by CSIRO Publishing. Cur¬
rent issues are freely available. The Proceed¬
ings are only available in soft copy. Articles
back to 1855 are available on-line.
The Papers & Proceedings of the Royal Soci¬
ety of Tasmania are freely available on- line,
but not for the last two years. There are,
apparently, no hard copies printed. Only
members of the Society who pay to do
so have access to the last two years of the
Papers.
The Proceedings of the Royal Society of
Queensland are apparently freely available
on-line. The Proceedings publishes only a
single issue a year, and then only in soft
copy.
The Journal of the Royal Society of Western
Australia is (since December 2015) no longer
printed, but is available only on-line to fully
paid members of the Society and approved
exchange partners and educational institu¬
tions.
The Journal of the Royal Society of New Zea¬
land is published quarterly in soft and hard
issues by Taylor & Francis. It has a 5-year
impact factor of 0.9 1 8. The current issue is
available on-line, but archive (permanent)
access to soft and hard copy costs AU$640 a
year. Earlier issues (1868 to 1961) are freely
available on-line.
What is to be done? Sixty years ago
Ronald Nyholm’s suggestion was not par¬
ticularly effective. And today it is even less
likely to succeed. I encourage review articles
and articles taking an historical approach to
the development of science and social phe¬
nomena. I also urge readers to consider writ¬
ing longer book reviews of recent books that
raise issues of interest, particularly of science
policy and history. This fits, I hope, with the
Society’s push to widen its membership from
the hard sciences to the social sciences and
to the arts and humanities. Please consider
the Journal for your next such paper.
3
Journal & Proceedings of the Royal Society of New South ^^es
Marks Editorial
Finally, Fd like to thank Ed Hibbert, Don
Hector, and Jason Antony for their assistance
in processing the Journals text.
30 November 2016
References
Nyholm, Ronald S. Presidential Address,
JProcRSNSW 89: N29, 1955.
http://biodiversitylibrary.org/ page/46 1 93345
4
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 5-16. ISSN 0035-9173/16/010005-12
Presidential Address
Donald C*A* Hector
The Royal Society of NSW
Email: dchector@royalsoc.org,au
Abstract
Donald Hector AM was President of the Royal Society of NSW from 2012 to 2016. In an address
marking the conclusion of his presidency, immediately following the annual general meeting of the
Society on Wednesday, 6 April 2016, he considered the nature of the complex problems that face
2 lst~century Australia, the way in which people tend to approach these highly-complex socio-techno
problems and the cognitive and cultural limitations they have in identifying solutions. In particular,
he considered the role that the Royal Society of NSW might play as it is re-established as a leader in
the intellectual life of NSW and of the country.
For many years, it was a practice of the
Society for the President to deliver an
address at the conclusion of the presidential
term. This custom fell into disuse in recent
years but with the change to the rules and
bylaws last year, it was decided to reintro¬
duce it.
My aim tonight is not to reflect on the
activities of the Society in the last four
years, other than in passing--- rather, it is
to attempt to chart a way for the Society as
it re-establishes itself as an intellectual force
in New South Wales and the country So I
shall limit my comments on the recent his¬
tory to these.
By far the most successfiil development
in the last several years has been the estab¬
lishment of the category of Fellow and the
elevation of the former Fellows to Distin¬
guished Fellowship. This raised considerable
interest in the activities of the Society and
we were fortunate that very capable people
indeed have accepted the invitation to Fel¬
lowship and that some of them have become
involved in the activities of the Society and
its governance. We expect that this will con¬
tinue and that we will see sustained growth
in all membership categories. But this will
only be the case if the activities of the Soci¬
ety are considered to be making a valuable
contribution to the public discourse. How
might we do this?
In September last year, for the first time,
the Society organised a forum with the four
Australian learned Academies. One of the
outcomes of the meeting was a list of major
challenges and issues where the Society
could contribute, taking a transdisciplinary
approach across art, science, literature and
philosophy. Of the issues identified at the
forum, one common characteristic they
shared was that they are all highly-complex,
socio-techno-economic problems. Most of
these are not limited to NSW nor to Aus¬
tralia— in many cases, they are global issues.
I would like to spend the remainder of my
address exploring how these complex prob¬
lems have come to be, why we see them in
the way we do and what we can do to con¬
tribute to a solution. I will take a historical
perspective and consider some issues around
philosophy and cognitive psychology that I
believe are important in framing these prob¬
lems and identifying solutions.
The way in which all animals interact and
survive in their environment is through solv-
5
Journal & Proceedings of the Royal Society of New South ’^^es
Hector— Presidential Address
ing problems. Humans have developed a
remarkable capacity for intellectualising
problems and solving them in the abstract.
Some of these problems can be simply stated
and have simple solutions — for example,
will I catch the bus to work this morning
or will I ride a bicycle? At the other end
of the spectrum, there are many problems
that can be both difficult to articulate and
to resolve. Contemporary examples of these
are: what are we to do about climate change?
or how can we provide a cost-effective health
system? I would like to briefly explore the
nature of problems and why some of them
are so difficult to understand and to solve.
The way in which we define and attempt
to solve problems today has its origins in
the philosophy of ancient Greece. Indeed,
the rediscovery of classical philosophy in the
13th and I4th centuries was a major influ¬
ence on the Renaissance. Let me refer to
an example. Many of you will have seen
this painting or be familiar with it. It was
painted by Raphael in 1509 and is a fresco
in the Apostolic Palace in the Vatican. It is
widely considered as one of the finest pieces
of art from the Renaissance, It is usually
referred to as The School of Athens (although
its formal name is Knowledge of Causes).
The School of Athens
The two central figures are Plato and Aris¬
totle but other Greek philosophers (Socra¬
tes and Diogenes) are also represented, as
are other philosophical influences from the
pre-Christian era. I will refer to this paint¬
ing again later to make some other points
but what the painting shows is the influ¬
ence of philosophy in Renaissance thinking.
It is intended to represent natural truth as
acquired through reason, arithmetic, geom¬
etry, astronomy, rhetoric and dialectic and
also represents art, music and poetry.
On the opposing wall is a second paint¬
ing, also by Raphael, called Disputation over
6
Journal & Proceedings of the Royal Society of New South Wales
Hector— Presidential Address
the Most Holy Sacrament. It was painted the
following year and shows God the Father
looking down on the resurrected Christ who
is flanked by the Virgin Mary and John the
Baptist and prophets and saints of the old
and new Testaments. Beneath are Popes,
saints and the faithful masses and with Aris¬
totle, head slightly bowed and his books on
the ground.
Disputation over the Most Holy Sacrament
On the one side of the chamber, is a
representation of knowledge and reason;
on the other, the realm of God. In many
respects, these two juxtaposed paintings
represent the thinking and belief-system of
that era and upon which the Renaissance
developed. Art can give great insight into
human thought — ^let us explore this notion
a little further.
Consider this example of ancient Greek
art. It is from a piece of pottery of an uncer¬
tain date and is thought to represent Euripi¬
des’ Medea. Its composition is what Paul
A piece of Greek pottery of an uncertain date
and is thought to represent Euripides Medea.
7
Journal & Proceedings of the Royal Society of New South Wales
Hector— Presidential Address
Feyerabend (1975) refers to as a “paratactic
aggregate”^ — ^ specially-structured group
of individual elements. Feyerabend sug¬
gested that this shows that Greek thought
was elemental in nature — they believed that
that everything in the world consisted of
atoms that were aggregated into bigger and
bigger things.
The story is told by the relationship of the
elements in the artwork. Another charac¬
teristic of Greek art was that it had no per¬
spective. Together, this suggests that Greek
thought was not developed into an integra¬
tive, representation of the world and that
the sense of perspective that is important in
modern representations simply had not yet
developed. This notion is reinforced by the
nature of the ancient Greek language. It is
also paratactic — it depends heavily on struc¬
ture. Their thinking — -the paradigms they
used — -were, perhaps, elemental, mechanis¬
tic and lacking in perceptual depth.
Now let us move to the very early Renais¬
sance — -about 1350. Consider the fresco
in Campo Santo, Pisa, by Francesco Traini,
called Triumph of Death.
Fresco in Campo Santo, Pisa, by Francesco Traini, Triumph of Death.
There is much similarity between the style
of Greek art and this painting: there is no
perspective and the story is told by structure
of the elements represented in the painting.
Let us now go forward about 100 years to
1430^ — this is a scene painted by Paolo
Uccello representing Mary approaching
a temple. In the space of 100 years or so,
perspective has started to emerge, giving
a sense of depth and three-dimensionality.
The style is more integrative, with the charac¬
ters becoming part of the scene, rather than
simply arranged in it.
8
Journal & Proceedings of the Royal Society of New South Wales
Hector— Presidential Address
A scene by Paolo Uccello representing Mary
approaching a temple.
Now let us return to our painting by Raphael,
The School of Athens. Painted 70 years later,
in 1509, and at the height of the Renaissance,
the style is entirely different. It is much more
fully developed: perspective is clear— it is a
representation of a three-dimensional scene
that truly appears to be in three dimensions.
Everything is integrated: the various philoso¬
phers and thinkers are engaged in conversa¬
tion with each other (even though some are
from different eras). It tells a story.
What I have tried to show here are some of
the foundational influences on the Western
way of thought, as represented through its
art. Whether or not Feyerabend’s theory is
correct is open to discussion but it is hard
to accept as coincidental the extraordinary
development in the sophistication of artistic
representation that happened at the same
time as the development of philosophical
thought in centres such as Florence and
Padua. As the influence of the Renaissance
moved from Italy across Europe, the centre
of intellectual thought gravitated towards
Holland, to (what is now) Germany and
to England. The discoveries and thinking
of Copernicus, Galileo, Bacon, Locke and
Newton, based on Greek philosophy, contin¬
ued to develop within the mechanistic Greek
paradigm — the universe was like a great
machine overseen by God. This thinking
prevailed until the 1 8th century when phi¬
losophers such as Kant and Hegel brought
different perspectives to our interpretation
of reality and the “interconnectedness” of
everything in the universe. Nonetheless,
the mechanistic paradigm persisted until
the late 1 9th century. At about this time,
biology and ecology began to develop and
the mechanistic paradigm was insufficient to
explain many of the phenomena that were
now being observed. A new model emerged
for explaining these — systems theory.
Mechanisms, like clocks, behave lin¬
early — -a disturbance to the mechanism
produces an effect in proportion to the
disturbance. The analytical technique
developed in Padua in the mid- Renaissance
works very well. If you have a problem, dis¬
assemble it into its component parts, solve
the component problems and synthesise a
solution to the original problem from these.
But systems do not work this way. They
are non-linear — a tiny disturbance in one
part of the system can result in a surprisingly
large disturbance in another. Systems can
appear to be stable but a small disturbance
can introduce major instability— they can
flip. They are characterised by subsystems
whose behaviour interacts with other sub¬
systems to influence the behaviour of the
whole — -you cannot predict the outcome
by simply adding the subsystem responses
together. At the heart of systems theory is
that everything in the universe influences
everything else. Systems theory and its
underlying philosophy of interconnected¬
ness and uncertainty was remarkably success¬
ful as a means to understand an enormous
array of phenomena from the behaviour
of ecosystems, to quantum mechanics, to
biological systems to the behaviour of high¬
speed aircraft and the control of equipment
9
Journal & Proceedings of the Royal Society of New South ^W^es
Hector — Presidential Address
in power stations and other industrial proc¬
esses.
After the Second World War, there was
massive rebuilding and restructuring of soci¬
ety. New problems started to emerge: how
to provide healthcare, establishing effective
educational systems, trying to make sense
of economics to avoid catastrophes such as
the Great Depression and deal with ever-
increasing environmental damage. Advanced
mathematical techniques such as linear pro¬
gramming, game theory, queueing theory,
marginal analysis and information theory
were developed as a consequence of milita¬
risation, in particular strategic analysis as the
Cold War deepened. But these were gener¬
ally unsuccessful in solving this new class of
problem. They were variously described as
“messes” and “wicked problems”. Generally,
there was no shortage of data to draw upon
to characterise the problem. But the systems
nature of these problems were surprisingly
resistant to analytical approaches. Typically,
these problems had social dimensions that
were difficult model.
Summarising, over the last 500-600 years
the way in which problems are identified
in characterised has evolved substantially.
We now think of problems in three broad
types:
But there is another dimension to problem¬
solving: the “domain of interests”: that is,
the people or stakeholder groups involved in
Problem type
Characteristics
Simple Problems (or mechanistic
or scientific problem)
Problems that can be represented using a mechanistic
model and resolved using the reductionist approach.
Single-Dimensional Complex
Problems (or technical or systems
problem)
Problems — ^ often of a technological nature — ^that can
be represented on one problem dimension.
Multi-Dimensional Techno-
Societal Problems
Problems that can only be represented on multiple
dimensions, considering issues such as moral status,
intrinsic character, value, beliefs, aesthetics etc.
Table 1: Increasing problem complexity
either the problem itself or any solution that
might be identified. The domain of interests
can range from a single individual trying to
solve a simple problem up to highly complex,
global problems whose domain of interest
extends across species and ecosystems. One
might conceive of three broad domains of
interest: unitary; pluralist; and disparate.
A unitary domain exists where there is
a single decision-maker or, if there is more
than one individual, where the decision¬
makers have a shared worldview and an
agreed determination in resolving the
problem. A pluralist domain is one where
there is a shared determination to problem
resolution but there are differing worldviews
among the stakeholders. Issues of power
and coercion are either explicitly or implic¬
itly set aside. And a disparate domain of
interests is where there are major differences
in underlying beliefs and values among the
stakeholders. The worldviews represented in
the domain may be in open conflict. There
may not even be agreement that a prob¬
lem exists or that action needs to be taken.
There may be distrust among stakeholders
10
Journal & Proceedings of the Royal Society of New South Wales
Hector— = Presidential Address
and there may be deliberate use of power to
coerce or frustrate problem definition and
decision-making. These three domains are
represented in table 2.
of problem can be identified as shown in
diagram 1:
Typel : these are problems which normally
yield to reductionist or systems-analysis
problem-solving approach. Traditional
scientific and engineering methodologies
can be applied such as mathematical mod¬
elling and computer simulation.
Type 2: these problems which due to their
complexity and systems nature require
a combination of reductionist, analyti¬
cal and hard- and soft-systems analysis
approaches.
Type 3: these problems are often unique and
always highly complex — this precludes or
severely limits the use of traditional sci¬
entific, engineering and systems analysis
approaches. Human stakeholders hold
Taking these two dimensions of problem
structure together, three fundamental types
apparently irreconcilable differences in
beliefs and values and are more than will¬
ing to exploit power imbalances coercively
to achieve their own ends. Moral status
of stakeholders and their interests may be
difficult to identify and some (for exam¬
ple, non-human species) may not be for¬
mally represented in the decision-making
domain.
This characterisation of Type 3 problems
that has emerged over the last half century
or so is not unique nor is it particularly
new. These “wicked problems” or “messes”
and have occupied the thoughts of opera¬
tional researchers for many years but with
only limited success. But what is diflFerent
here is the representation of these problems
on two dimensions — recognising that the
influence of differing worldviews and the
Domain of
INTERESTS
Characteristics
Unitary
A single decision-maker or a group of decision-makers and other stake¬
holders which have the same interests and similar worldviews.
Pluralist
Decision-making interests are largely aligned but there may be many
different worldviews among stakeholders. However, they share the same
determination or interest in arriving at a satisfactory problem resolution.
Power is equally shared among constituents or, because of the shared
determination to resolve the problem, issues of power are set aside.
Disparate
There are major differences in underlying beliefs and values and the
interests of stakeholders may differ widely. There may be a lack of shared
determination to resolve the problem, distrust of the motives and inten¬
tions of other interests, and even specific intention not to see the situation
resolved and to derail attempts to agree upon the problem definition or
eflForts to proceed. There may also be significant power imbalances among
the constituents and these are used coercively.
Table 2: A further dimension of problem complexity —the Domain of Interests
11
Journal & Proceedings of the Royal Society of New South
Hector “—Presidential Address
coercive use of power has enormous influ¬
ence both on characterising the problem
and moving towards some resolution. An
important point to note is that describing or
structuring the problem is fundamentally a
human, social construct. Worldviews and
belief-systems are at the heart of both defin¬
ing and solving problems.
Let me summarise my argument so far.
As the humans have evolved, so too has
the way in which we conceive of and attempt
to solve problems. The worldview of the
Domain of interests
Diagram 1 —problem taxonomy.
ancient Greeks persisted for well over a mil¬
lennium. But in just a couple of hundred
years, the Renaissance brought a dramatic
change in the Western world— “there was
a flourishing of thought and a number of
different philosophical approaches emerged.
But as population and social complexity
increased, so too did the complexity of the
problems that confront us. Not only are
the technical aspects of the problems chal¬
lenging but they are further complicated by
the divergence in worldviews that occurred.
In the last century or so, social and cultural
influences in an increasingly liberal soci¬
ety have added another dimension to the
way in which we need to consider complex
problems. The second dimension of prob¬
lem structure that I have proposed here is
predominantly about influence and power
and is a major obstacle in solving the highly
complex socio-economic problems.
Before exploring how these types of prob¬
lems might be addressed, I would like to take
a brief diversion to outline the philosophi¬
cal framework upon which Anglo-American
society (by which I mean the various coun¬
tries around the world that emerged from
or were strongly influenced by Britain) has
developed in the period since the late I6th
century. I confine my remarks to the Anglo-
American philosophical framework because
Royal Societies around the world are crea-
12
Journal & Proceedings of the Royal Society of New South Wales
Hector—" Presidential Address
tures of this. It has also been very influential
in the development of the modern era. Time
does not permit a detailed examination of
these, so I will just outline them briefly.
What was originally called “philosophy”
and has evolved into scientific enquiry is
dependent on the scientific method of con¬
jecture and refutation. It is founded on a
rationalist philosophy and traces its origins to
thinkers such as Bacon, Locke and Newton.
One of the principles of rationalism is that
we can acquire knowledge in two ways: one
is empiricist (interpreting knowledge that we
acquire through our senses); and the other
is intuitive and deductive (there is some
knowledge that we can acquire through
thought and deduction alone — -mathemat¬
ics being an example). It is oversimplifying
somewhat to say that scientific enquiry is
entirely rationalist— there are many factors
that influence it, including sociological ones.
But its intention is to converge upon some
notion of truth through rigorous, intellec¬
tual enquiry.
Political and social institutions are gener¬
ally framed on different philosophical prin¬
ciples - — they are utilitarian. Utilitarianism
originates with Bentham and Mill and has
been developed by many others. Its original
concept was that a “good” act is one that
maximises pleasure. Unlike rationalism, it
is less concerned about finding truth; rather
it is a normative ethical system— it attempts
to define a set of rules for society to live
by. Over the last couple of hundred years,
“pleasure” has been replaced with “benefit” or,
more recently, “happiness”. Most economic
analysis is utilitarian in its nature: what will
deliver the maximum benefit for the mini¬
mum cost? Similarly, our political systems
attempt to arrive at maximising public good
(or happiness) with minimum interference
with individual liberty — they are funda¬
mentally utilitarian, liberal frameworks.
Legal institutions are different again.
They are also based on normative philo¬
sophical principles but are framed around
deontological or duty-based ethics. One
particularly influential philosopher in this
area was Kant who argued that a “good” act
is one in which one does ones duty. Duty
can be defined in terms of a legal code or
duties that emanate from moral good. Kan¬
tian ethics is controversial but nonetheless
the British legal system is largely duty-based
system. (For example, sections 180 to 183
of the Corporations Act defines duties that
must be observed by company directors.)
So, the society in which we find ourselves
today is largely the product of three philo¬
sophical systems that are becoming ever¬
more influential and, in many aspects, are
replacing the influence of religion that until
relatively recently dominated our value-sys¬
tems. The extent to which belief (whether
religious or humanist or some other value-
based system) influences decision-making is
of critical importance in solving the highly-
complex Type 3 problems that prove so chal-
lenging.
I will now briefly explore some cognitive
psychology in an attempt to identify the way
in which these Type 3 problems might be
addressed. There is a large body of literature
in cognitive psychology relating to problem¬
solving originating, in the 1920s and 1930s.
I will confine myself just to drawing a few
points from this literature.
One of the key researchers in this area
was Hammond (1955) who integrated the
work of a number of eminent psychologists
relating to the way in which people respond
to cues that they receive. Researchers found
that people form judgements and make
inferences based on observations that are
weighted according to their experience and
other subjective influences. The analogy of
the “lens model” was created ^ — just as light
13
Journal & Proceedings of the Royal Society of New South Wales
Hector— Presidential Address
is distorted by an optical lens, giving differ¬
ent images to different observers depend¬
ing on their position, so too do individu¬
als involved in a complex situation form
different perceptions of the problem and
the path forward. Hence, there can be no
objectively-determined understanding of
complex problems.
In a ground-breaking piece of work in
the 1950s, Miller (1955) found that people
have a very limited capacity to retain pieces
of information in their minds (somewhere
between five and eight pieces of information
at any one time) but an extraordinary capac¬
ity to recall information to mind to proc¬
ess it. Another related body of research by
Boulding found that people form “images”
or mental representations of situations that
are important in the way in which they reach
decisions. They imagine what the future
might be and then strategise to achieve it.
These images are not simply mental pictures,
rather they are complex mental representa¬
tions of situations that we are attempting
to understand. An interesting example of
this is the “cognitive map” that we form to
help us relate to our situation in the physical
world— not only is it a locational map, it is
a representation of self and our relationship
with the physical world. This fundamen¬
tal cognitive process probably underlies the
human penchant for representing complex
information in a wide range of graphical and
visual formats, such as maps.
Many of these mental phenomena are
not specific to humans — all cognisant
animals seem to utilise them. It is how
cognisant beings deal with the enormous
complexity of the world in which they find
themselves. The complexity is too great to
comprehend, so a form of thinking — intui¬
tive thought — evolved to make sense of
it. At some point our evolution, humans
developed the capacity for rational think¬
ing. It is the capacity for rational thought
that makes humans sapient (some other
animals appear to possess limited capacity
for rational thought but there is none that
comes close to humans). But the capacity
for rational thought is bounded — the world
is far too complex for the human mind to
comprehend it completely.
On one hand, intuitive thought is used by
all cognisant animals. It is instinctive and
quick and the main mechanism by which we
survive. On the other, rational thought is
largely peculiar to humans. It is slow, delib¬
erate and it is learnt. In the 1970s and 1980s,
work byTversky and Kahneman (1974) and
others found that intuitive thought is subject
to a range of biases and that these have a
significant impact on the success of decision¬
making. Rational thought (or least some of
the means to it) can be taught and improved
but it is error-prone. Whereas intuition is
subject to bias, rational thought is subject
to error.
But we need to put these mental represen¬
tations and processes into a both a chrono¬
logical and cultural context. This requires
another cognitive device- — -the narrative.
Narrative and story-telling is as old as
humanity itself It predates writing and
occurs in every human society and culture.
Throughout most of history, story-telling has
been the principal means by which knowl¬
edge is transferred from one generation to
the next. There are various theories of nar¬
rative but they share some common char¬
acteristics. They are always about people
or things and a group of characters forms
part of the thread that holds the narrative
together. They are developed against an
explicit set of values or a moral standard
against which the actions in the narrative
can be evaluated. Until the 1970s, narrative
was thought to be simply a cultural artefact,
but now it is considered to be a fundamen-
14
Journal & Proceedings of the Royal Society of New South Wales
Hector — Presidential Address
tal cognitive process. While cognitive maps
provide the three-dimensional framework
that we use to relate to the real world, nar¬
rative adds the fourth dimension — ^time. It
also provides the means to fill in the gaps in
our understanding and to make our mental
representation coherent with our experience
and our worldview. In other words, we make
things up — we confabulate— -to fill in the
gaps in our knowledge and most importantly,
in order to make our representation of the
problem conform with our belief-system.
So, let me summarise these few fragments
of psychology. No two individuals see a
problem in exactly the same way— we are
all looking at things through “lenses” that
distort our view of reality according to
our perceptions and experience. We form
images of problem situations that are heav¬
ily influenced by our philosophical frame¬
work and belief-system. Our immediate
response to problems is intuitive but this
is subject to bias. A more measured ana¬
lytical approach— rational thought— can
be learnt but we must remain aware that
we can make mistakes. These two thought
processes have been described as two dif¬
ferent systems but that misunderstands the
fundamental nature of cognition — they
are a single system responding to diflPerent
stimuli and this system exhibits all the non¬
linear and unexpected characteristics that
one would expect. In order to make sense of
the enormous complexity we encounter, we
confabulate to make sense of things that we
do not understand to make them conform
to our notions of reality.
So how might we move forward?
Recognising the enormous human crea¬
tivity available to us through combining our
capacity for intuitive and rational thought,
we can use the enormous body of knowledge
(that continues to grow at an exponentially
rate) and our capacity for rational analysis
to gain much greater insight into problems
that were previously unassailable. We can
imagine what futures might look like. If
we remain conscious of the bias associated
with intuition and alert to the ever-present
chance of error with rational thought, we
can reduce the chance of serious mistakes.
Recognising the systems nature of cognition,
we can harness both intuitive and rational
thought to bring great creativity. Because
we can recognise that various stakeholders
in situations will approach the problem from
different perspectives, we can accept this as
fundamental to the human condition and
that should facilitate understanding. The
big challenge is to embrace the complexity
of the problem— particularly the sociologi¬
cal dimensions— to overcome the inherent
bias that we all hold to find common ground,
rather than focus on the differences.
Most importantly, we can write narratives.
Drawing upon our diverse experience, these
narratives can engage people with a wide
range of worldviews and draw them along
with us.
And now I come to my final point. The
Royal Society of NSW is uniquely placed
to provide leadership in this type of com¬
plex analysis. The wisdom of the founders
in defining such a broad remit of human
knowledge — science, art, literature and
philosophy — was truly prescient and rec¬
ognised the ever-increasing complexity of
modern life. But we need to change if we
are to maximise our impact. Historically,
the Society has focused on the sciences: in
its early days, the physical sciences— physics,
chemistry and geology- — ^and, later, zool¬
ogy, botany and biology. Only recently, have
we extended into the other areas of human
knowledge encompassed by our charter. We
need to attract Fellows and Members from
all fields of human knowledge, if we are to
engage in the representation and solution
15
Journal & Proceedings of the Royal Society of New South Wales
Hector — Presidential Address
of Type 3 problems. We need more writers,
artists, sociologists, musicians and historians.
Only then, will we be able to completely
engage with the community. That is not to
say that we should abandon our scientific
heritage — quite the opposite, most of the
problems that the world faces today have
enormous technological challenges. But
these solutions will not be found in science
and technology alone — they will require the
engagement of non-scientists in terms they
can understand.
References
Feyerabend, R {\37 5), Against Method, (3rd
edn., 1993), Verso, London, pp 170-187.
Hammond, K.R., (1955), Probabilistic
Functioning and the Clinical Method,
Psychological Review, vol. 61, no. 4, 1955,
pp 255-262.
Miller, G.A., (1956), The magical number
seven, plus or minus two: some limits on
our capacity for processing information, The
Psychological Review, 63, 2, pp 81-97.
Tversky, A., Kahneman, D. (1974), Judgment
under uncertainty: heuristics and biases,
Science, New Series, 185, 4157, pp 1 124-
1131.
General bibliography
Ackoff, R.L., (1979), The future of operational
research is past, Journal of the Operational
Research Society, 30, 2, pp 93-104.
Gabriel, Y, (1991), Turning facts into
stories and stories into facts: hermeneutic
exploration of organisational folklore,
Human Relations, 44, 8, pp 71 1-724.
Hammond, K.R. (2007), Beyond rationality:
the search for wisdom in a troubled time,
Oxford University Press, New York, USA
Hector, D.C., Petrie, J. et at, (2009), A
problem-structuring method for complex
societal decisions: its philosophical and
psychological dimensions, European Journal
of Operational Research, 193, 3, pp 693-708,
doi:10.10l6/j.ejor.2007.06.058.
Kintsch, W, van Dijk, TA. (1978), Toward a
model of text comprehension and production.
Psychological Review, 85, 5, pp 363-394.
Kubovy, M. (1986) The psychology of
perspective and Renaissance art. Cambridge
University Press, Cambridge, UK.
Miller, K.D., Waller, H.G. (2003), Scenarios,
real options and integrated risk management.
Long Range Planning,. 36, pp 93-107.
Pentland, B.T., (1999), Building Process
Theory with Narrative: From Description to
Explanation, Academy of Management Review,
24, 4, pp 71 1-724.
Randall, J.H., (1940), The development of
scientific method in the School of Padua,
Journal of the History of Ideas, 1, 2, pp 177-
206.
Rosenhead, J. (ed.), (1989), Rational analysis
for a problematic world: problem structuring
methods for complexity, uncertainty and conflict,
John Wiley & Sons, Chichester, UK
Russell, B. (1946), History of Western
Philosophy: and its connection with political
and social circumstances from the earliest times
the present-day, (2nd edn., 1961), Routledge,
London.
Tolman, E.C., (1948), Cognitive maps in
rats and men. Psychological Review, 55, 4, pp
189-208.
von Bertalanffy, L., (1950), An outline of
General System Theory, British Journal for the
Philosophy of Science, 1, 2, pp 134-165.
Donald Hector AM was President of the Society from 2012 to 2016.
16
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 17-33. ISSN 0035-9173/16/010017-17
Deep-earth methane and mantle dynamics:
insights from northern Israel, southern Tibet and
Kamchatka
William L. Griffin^*, Sarah E.M. Gain^ David T. Adams^, Vered Toledo25
Norman J. Pearsoni and Suzanne Y. O’ReiUyi
^ ARC Centre of Excellence for Core to Crust Fluid Systems, EPS, Macquarie University,
NSW 2109, Australia
^Shefa Yamim (A.T.M.) Ltd., Akko, Israel
* Corresponding author.
Email: bilLgriffin@mq.edu.au
Abstract
The oxidation state of fluids in Earth’s mantle affects processes ranging from volcanism and the forma¬
tion of the crust, to the generation of many types of ore deposits. In general, the lowest oxidation state
of the mantle (and hence its fluids) is defined by the buffer reaction 2FeO 2Fe + 02. However,
unusual mineral assemblages that require far more reducing conditions are found in volcanic rocks
from a variety of geotectonic settings, raising questions about how such conditions can be generated
in the mantle. Examples from northern Israel, Tibet and Kamchatka suggest that interaction between
magmas and methane-hydrogen fluids derived from the deep Earth have generated highly reducing
conditions within some volcanic plumbing systems. Such systems appear to be related to the margins of
tectonic plates, including zones of continent-continent collision and/or deep oceanic subduction, and
transform faults extending deep (up to 200 km) into the Earth’s mantle. This represents an important
but previously unrecognized fluid-transfer process within the mantle.
Foreword
his paper is both a condensation and an
expansion of the 2015 Clarke Memo¬
rial Lecture, given by WLG at Macquarie
University in August 2015, which summa¬
rized recent work by the ARC Centre of
Excellence for Core to Crust Fluid Systems
(CCFS). We hope to provide both geolo¬
gists and non-specialists with a glimpse into
some recent exciting developments in Earth
Science, and to show how the integration of
observations at scales from microns to moun¬
tain ranges can give us a new picture of how
Earth works. We are trying to understand
processes not previously recognized, and the
paper therefore contains some “interesting”
speculations, which we hope can generate
(polite) discussion.
Introduction
The nature of the rocks in Earths deep
mantle (from the base of the crust to the top
of the core; Fig. 1), and clues to their forma¬
tion, are generally hidden beneath our feet
in the vast space of inner Earth. However,
small samples of the mantle are brought to
the surface by some volcanic eruptions in
which the magmas originated at depths of
250—100 km. Such magmas can rise to the
surface at speeds of 1-4 km/hour, entraining
fragments of deep-seated rocks and minerals
(xenoliths and xenocrysts; O’Reilly and Grif¬
fin, 2010). Larger samples of mantle rocks,
tens to thousands of cubic km in volume,
can be brought to the surface by geodynamic
forces in regions where tectonic plates collide
(e.g., Tibet, the Andes, New Guinea).
17
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
Figure 1 . (a) Cross-section of Earth s interior, showing the divisions defined by mineralogy; (b)
Cross-section emphasizing the convection of the mantle, driven by the heat in the core; (c) cartoon
of deep-seated mantle plumes, in this case producing oceanic islands like Hawaii; smaller upper-
mantle plumes may rise from the Transition Zone and produce much less magma.
This report focuses on three locations that
have yielded surprising discoveries about the
nature of some mantle rocks and minerals
and their conditions of formation deep in
the mantle. Material from two of these areas
(S. Tibet, N. Israel) is being studied by the
CCFS team, while recent discoveries in the
Kamchatka Peninsula of NE Siberia provide
important comparisons.
The Deep Earth and the Importance of
Fluids
The large-scale structure and composition
of Earth’s convecting mantle (the astheno-
sphere), extending from 100-200 km below
Earth’s surface to the outer edge of the core
at -2,900 km, are generally understood (Fig.
1). The uppermost (lithospheric) mantle is
relatively cool and coupled to the crust; (10
to 40 km thick in oceanic and old conti¬
nental regions respectively); these make up
18
Journal & Proceedings of the Royal Society of New South ^X^ES
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
the lithosphere, and Earth’s tectonic plates.
The mantle is made up largely of ultramafic
rocks— -high in MgO and low in Si02. At
depths between 410-660 km, in the Tran¬
sition Zone, the dominant minerals (oliv¬
ine, pyroxenes, garnet) go through a series
of changes (phase transitions) to denser,
higher-pressure forms, which dominate the
mineralogy of the lower mantle (>660 km).
The convection of the asthenosphere drives
(or responds to) the movement of the tec¬
tonic plates that make up Earth’s crust (Fig.
lb). In the upwelling parts of the convec¬
tion system, decompression induces melting,
with the production of a range of magma
types, at relatively shallow levels of the
asthenosphere. Where plates descend into
the mantle at subduction zones, the intro¬
duction of water helps to cause large-scale
melting, producing volcanic arcs^ — ^ ranges
of volcanoes like the Andes. Whether the
whole mantle convects in the same way is
still debated, but it is clear that material
does emerge from the Transition Zone or
even the core-mantle boundary, in the form
of plumes, which bring hot mantle to near
the surface. The results are visible in chains
of volcanoes such as Hawaii; these '‘hotspot
trails” reflect the movement of an oceanic
plate over a stationary plume (Fig. Ic).
The role of fluids in all of these mantle
processes is a major focus of the research
program in the ARC Centre of Excellence
for Core to Crust Fluid Systems (CCFS).
The presence of fluids helps to determine
the viscosity of the convecting mantle, the
temperatures at which it melts to produce
magmas (which also are fluids), the nature
of volcanic eruptions (explosive vs quies¬
cent) and the compositions of the magmatic
products. We need to know the nature and
distribution of different types of fluids in
the mantle if we are to understand these
processes.
Oxidation-reduction (redox) reactions, in
which oxygen is transferred from one min¬
eral to another, or from a fluid to a rock or
vice versa, play a vital role in many Earth
processes, especially in the genesis of igneous
rocks, and the movement and reaction of
fluids at all- levels of Earth’s internal archi¬
tecture. Redox reactions, as well as pres¬
sure (P) and temperature (T), can control
which minerals are stable in rocks, and the
distribution of water, carbon dioxide (CO2),
methane (CH4), hydrogen and other compo¬
nents in fluids moving through the mantle
and crust. Via such fluids, redox reactions
also are very important in the formation of
many types of ore deposits.
The “redox state” of a chemical system can
be expressed in terms of the oxygen fugacity
(/O2), which is simply the partial pressure of
oxygen in an ideal gas of a given composition.
It commonly is expressed as values relative to
the y02 of a known reaction (a “buffer”; Fig.
2). For example, a relatively high oxygen
fugacity (“oxidizing” conditions) would be
defined by the reaction:
Fayalite (Fe-olivine) + 02^ magnetite +
qtz
Fe2Si04+ O2 Fe304 + Si02 (1)
the buffer
A much lower (more “reducing” condi¬
tions) would be defined by a reaction such
as:
Iron + oxygen wustite
2Fe + 02-^2Fe0 (2)
the buffer.
Most magmatic rocks in Earth’s crust
record jQ>2 around the FMQ buffer; the
upper mantle tends to have lower f^2^ ^.nd
to become generally more reducing (lower
19
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
jO^ with depth (Fig. 2). The IW buffer is
thought to represent a lower limit for j02
in the mantle, simply because mantle rocks
contain so much FeO that it would be hard
to reduce it all to Fe.
However, we do know that conditions
more reducing than the IW buffer must
occur locally in the mantle, partly because
the mineral moissanite (silicon carbide, SiC)
is found in some mantle-derived magmatic
rocks such as kimberlites (the host rock of
most diamonds) and even as rare inclusions
in diamonds. The stability of moissanite is
controlled by reactions such as:
Enstatite + carbon moissanite + forster-
ite
MgSi03 + C SiC + Mg2Si04 (3)
As we study other mantle-derived rocks, we
are finding that moissanite is unexpectedly
common in the mantle, and can be accompa¬
nied by a wide range of other “super-reduced”
minerals (e.g., metallic elements, carbides,
nitrides) that require even more reducing
conditions than moissanite itself. These
occurrences raise two important questions:
(1) what sort of processes can generate such
conditions in Earth’s mantle? and (2) why
didn’t these minerals react with the more oxi¬
dized mantle in which they are embedded?
Recent studies also have shown that
remarkably similar super-reduced mineral
assemblages can be found in mantle-derived
rocks from completely different tectonic set¬
tings. This suggests the widespread opera¬
tion of poorly-understood processes, not
previously recognized in the mantle. Can
the similarities and differences between
some of these occurrences provide clues to
the nature of those processes?
Here we will give brief descriptions of
three such occurrences, in southern Tibet,
northern Israel and the Kamchatka volca¬
noes of NE Russia; by examining their simi-
Figure 2. Oxidation-reduction (redox) reac¬
tions and their control on fluid compositions,
(a) Oxygen fugacity vs depth (at 1500 °C) for
some redox buffers; shaded field shows the
range of jOj measured in rock samples from
the upper (lithospheric) mantle, all above the
Iron-Wustite (Fe-FeO) buffer; (b) a crystal (4.1
mm long) of moissanite from the Mt. Carmel
area, Israel; (c) relative abundances of different
species in C-O-H fluids as a function of j02
(after Kadik, 1997). Below the Iron-Wustite
buffer the fluids are completely dominated by
CH4 and H2. EMOG/D, the buffer reaction
Enstatite + magnesite = olivine + graphite/
diamond.
20
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. Deep-Earth Methane and Mantle Dynamics
larities and differences, we can speculate on
a common process.
Study Areas
Tibet
In southern Tibet, the Yarlung-Zangbo
suture zone (Fig. 3) marks the great con¬
tinental collision between India and Asia,
which began ca 55 million years ago and
continues today, pushing up the Himalayan
Mountains. Scattered along this suture zone
is a line of peridotite massifs — - fragments of
Earth s mantle, up to 1 000 cubic kilometers
in volume. Some of these contain minable
bodies of chromite ore, which formed at
shallow depths (5-15 km) ca 325 million
years ago, when the peridotites were part of
the mantle above a subduction zone. How¬
ever, structures and relict minerals in the
chromite ores and the peridotites indicate
that these rocks were later subducted down
as far as the Transition Zone (>410 km),
where they remained for about 200 million
years (McGowan et al., 2015). During their
residence in the Transition Zone, the perido¬
tites probably heated up to 1400—1500 °C;
this heating, and their composition, would
result in them becoming buoyant relative to
the surrounding mantle.
The excavation of these peridotite bodies
from the Transition Zone back to the surface
can be attributed to the forces exerted by a
later slab, subducting during a plate collision
event and penetrating into the Transition
Zone. As the slab stalled, it began to roll
Figure 3. Bird’s-eye view (looking north) of the Tibetan Plateau, bounded in the south by the
Himalayas. Black lines show sutures between crustal blocks that have drifted together to form
the region. The southernmost line marks the Yarlung-Zangbo suture zone, where numerous
bodies of mantle peridotite (yellow) were emplaced 60-70 million years before the collision of the
Indian plate with Asia initiated the rise of the Himalayas. Many of these bodies contain diamonds,
super-reduced mineral associations and evidence of derivation from the Transition Zone; the red
arrow indicates the famous Luobusa peridotite. Map courtesy of Google Earth.
21
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
back, causing the mobile asthenospheric
mantle (Fig. 1, Fig. 4) to flow in to fill the
“gap”. Dynamic modeling (Afonso and Zlot-
nik, 2011; McGowan et al., 2015) has shown
that this can create a broad upwelling in the
deep mantle, near the top of the Transition
Zone. As slab rollback continues, this broad
upwelling rapidly develops into a narrower
channel of upward-flowing asthenosphere;
this upwelling combined with the compo¬
sitional buoyancy of the peridotite bodies
to bring them rapidly (in 6~8 million years)
from the Transition Zone to crustal levels,
where at least some became the seafloor in
small ocean basins (Liu et ak, 2015).
Small diamonds were first recovered from
some of the chromite ores in the peridot-
ites more than 40 years ago (Fig. 5; Bai et
al., 1993). Since then diamonds have been
extracted from both the ores and the perido-
tites in at least seven tectonically emplaced
mantle-derived bodies along the Yarlung-
Zangbo suture zone, in peridotites on older
sutures in farther north in Tibet (Fig. 3), and
in similar bodies in the Polar Ural Moun¬
tains of northern Russia (Yang et ak, 2014,
2015). These diamonds have created both
interest and disbelief, because they differ in
many respects from “normal” diamonds with
which most geologists are familiar.
“Normal” diamonds come from magmatic
bodies of kimberlite (an alkaline igneous
rock) that typically occur in the stable cra-
tonic areas of continents. Most diamonds in
the kimberlites are fragments from the con¬
tinental roots, and some may have resided
there for billions of years; they usually form
octahedral crystals, which may be rounded
by chemical resorption.
The Tibetan diamonds, in contrast,
strongly resemble synthetic diamonds that
are produced industrially in America, Russia
and China. They have smooth cubic faces,
which are rarely seen on kimberlitic dia-
Figure 4. A thermo-mechanical model (Afonso
and Zlotnik, 2011) showing how the rollback
of a subducting slab sets up forces that lead to
rapid upwelling of the mantle from the Transi¬
tion Zone. The blue marker represents a mass
of much older, more depleted peridotite rising
from the Transition Zone to the ocean floor.
(After McGowan et al., 2015).
22
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
monds (Fig. 5); they contain inclusions of
metallic alloys (Ni-Mn-Co); their carbon is
isotopically very light; they contain nitro¬
gen as single atoms, whereas nitrogen in
kimberlitic diamonds is largely recombined
into N-N pairs or N4 tetrahedra. However,
several diamonds have been found in situ
in the chromites (Yang et ah, 2007, 2014)
where they are surrounded by zones of amor¬
phous carbon. Detailed studies (Howell et ah,
2015) strongly suggest that the diamonds in
the peridotites and chromite ores are in fact
natural and represent a new environment for
diamond formation in the mantle-— but one
that is not well-understood.
Diamonds do not require strongly reduc¬
ing conditions to form in the mantle (EMOD
buffer. Fig. 2). However, the diamonds in
the peridotites of Tibet and the Polar Urals
are accompanied by a great variety of “super-
reduced” minerals (Yang et al., 2007, 2014,
2015). These include moissanite (silicon
carbide), native elements (Fe, Ni, Ti, Si, Cr,
Al), carbides of titanium and other elements,
silicides of iron and titanium, and titanium
nitrides. Many of the more unusual minerals
occur as inclusions in crystals of corundum
(aluminium oxide, AI2O3) with unusually
high contents of titanium (Xu et al., 2015).
Aside from the very low 7O2 required by
many of these minerals, it is difficult to pre¬
cisely define the conditions of their forma¬
tion. Many of the minerals imply high tem¬
peratures (I2OO--I5OO °C), and the inferred
presence of stishovite (a high-pressure form
of quartz) around some super-reduced phases
suggests depths >300 km (Dobrzhinetskaya
et al, 2009).
Northern Israel
In the Mount Carmel area of northern
Israel (Fig. 6), exploration for gemstones by
Shefa Yamim Ltd. has discovered a remark¬
able assemblage of super-reduced minerals.
Figure 5. Microdiamonds from Tibetan peri¬
dotites (a) SEM images of diamonds, with
octahedral and cubic faces; (b) diamond sepa¬
rated in CCFS labs, showing yellow colour
and smooth faces; (c) polished section showing
melt inclusions of Ni-Mn-Co alloy. Repro¬
duced from Griffin et al. (20 1 6) by permission
of Oxford University Press.
23
Journal & Proceedings of the Royal Society of New South ^VC^es
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
comparable in many ways to the Tibetan
assemblage (Table 1). These occur especially
in Cretaceous (ca 99-94 million year-old)
volcanic rocks on Mount Carmel, and in
alluvial deposits derived from them. The
volcanism occurs near the Dead Sea Trans¬
form Fault, which represents the boundary
between the Arabian plate and the northern
(Mediterranean) edge of the African plate.
However, the Cretaceous eruptions pre¬
date the first known movements (Miocene)
on the fault by about 70 million years (Sass,
1980; Garfunkel, 1989). Gas-rich explosive
eruptions produced thick pyroclastic depos¬
its (Fig. 6b) and some flows of frothy, glassy
lavas (Sass, 1980). The primary magmas are
mildly alkaline basalts. The presence of frag¬
ments of dense mantle rocks (garnet pyrox-
enites and websterites; Mittlefehldt, 1986;
Kaminchik, 2014; this work) shows that the
magmas erupted very rapidly once they had
risen to depths of around 80 km.
In addition to gem-quality sapphires,
rubies and “normal” diamonds, the Shefa
Yamim exploration program has produced
unusually large crystals of moissanite (Fig.
2) and large grains (up to 2.5 cm across)
Figure 6. (a) Geological map of northern Israel; Mt Carmel area is circled, and volcanic centres
are marked by red dots; (b) sawn section, ca 30 cm high, of volcanic tuff from Mt Carmel.
24
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
Figure 7. Fragment of non-gem corundum
from Mt. Carmel; dark irregular blobs are
pockets of melt trapped within and between
grains of transparent corundum. (After Griffin
et ak, 2016b).
of non-gem corundum” (Fig.7). The latter
has proved to be a Rosetta Stone for under¬
standing the development of highly reducing
conditions in Earth’s mantle.
Irradiation of the non-gem corundum
with electrons produces a bright pink fluo¬
rescence (cathodoluminescence, or CL; Fig.
8), which shows remarkable patterns that
outline the growth of the crystals. Micro¬
chemical analysis shows that variations in the
CL response correlate with differences in the
Ti content of the corundum — -light areas
fluoresce brightly, while high Ti contents
induce darker shades. The patterns show
that individual samples of the corundum
typically consist of many smaller crystals,
with pockets of melt trapped along the grain
boundaries and within the crystals (Fig. 7,
8). These melt pockets now consist of cal-
cium-aluminium-magnesium silicate glass
and minerals that crystallized from it before
the volcanic eruption carried the corundum
Figure 8. Cathodoluminescence images of
polished fragments of non-gem corundum.
Real colours range from light to dark pink to
nearly black. Light areas have low concentra¬
tions of Ti; dark areas are zoned toward melt
pockets and have up to 2.6 at.% Ti. (a) mul¬
tiple “stacked” crystals of corundum outlined
by high-Ti zones; (b) detail of crystal showing
growth zoning that outlines hollow (“hopper”)
faces and internal cavities filled with glass
(dark CL; frozen melts). (After Griffin et ak,
2016b).
25
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
fragments to the surface — so these minerals The corundum and its trapped minerals
were crystallizing in (or somewhere above) record very reducing conditions. Micro-
the magmatic plumbing system. chemical analysis shows that the Ti substi¬
tuting in the corundum is the highly reduced
Figure 9. Four types of melt pocket in non-gem corundum from Mt. Carmel, illustrating some
of the unusual, highly-reduced mineral associations, (a) SEM false-colour phase map showing
minerals and glass in a typical silicate melt pocket in corundum; (b) Ti map showing nitrides,
borides and silicides filling internal cavities in hopper crystal of corundum; (c) back-scattered
electron (BSE) image of TiC crystallized from an Fe-silicide melt, enclosed in corundum; (d)
BSE image of composite pocket of native vanadium, fluorite (CaF2) and hibonite (a Ca-Al oxide)
enclosed in corundum.
26
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
form Ti3+ (most Ti minerals contain the
more oxidized TP+), and one of the most
common minerals in the melt pockets is tis-
tarite (Ti203; Fig. 9a). Reduction ofTi02
(Ti4+) to Ti203 (Ti3+) requires ^"“8 orders
of magnitude below the IW buffer; these
are seriously reducing conditions. However,
even lower JO 2 is required by the presence of
TiO (with Ti2+) as a separate mineral, and as
a component in other oxides (Fig. 2). Other
low- j02 minerals in the melt pockets include
kharambaevite (titanium carbide, TiC), iron
silicide (Fe3Si), titanium nitrides, native iron
and native vanadium (Fig. 9b-d). The crys¬
tallization sequence of minerals in the melt
pockets suggests a continuous decrease in
j02 during their formation.
Moissanite has been found included in the
corundum, suggesting that both were part
of the same magmatic system. The CF pat¬
terns of the corundum show that it grew as
“hopper crystals” (Fig, 9), with hollow faces
and branching internal cavities. This type of
growth is characteristic of rapid crystalliza¬
tion from fluids that are supersaturated in a
component, in this case AI2O3. There is no
obvious genetic link between the diamonds
and the corundum, although exploration
has recovered a few microdiamonds similar
to those found in Tibet. However, the abun¬
dance of carbide minerals, and breccia veins
full of amorphous carbon cutting the grains
of corundum, imply the presence of fluid(s)
very rich in carbon. These two observations
may be the key to the origin of this bizarre
mineral system (see below).
It is difficult to constrain the temperature
and depth of the fluid plumbing system. As
in the Tibetan example, some of the minerals
present (and the presence of glass) suggest
temperatures in the range 1200-1500 °C.
The presence of the feldspar anorthite (CaAl-
251203) in some melt pockets suggests the
reaction corundum + melt anorthite,
which would constrain the depth to between
30 and 100 km. Fragments of mantle rocks
found in the volcanic ashes suggest that the
lithospheric mantle (Fig. 1) beneath the area
at the time of eruption was less than 100
km thick.
Kamchatka Peninsula, NE Russia
The Kamchatka peninsula (Fig. 10) hosts
many large, active strato volcanoes, reflect¬
ing the ongoing subduction of the Pacific
plate beneath NE Asia. From November
2012~September 2013, fissure eruptions
occurred on the flank of the Plosky Tol-
bachik volcano. The first eruptions, espe¬
cially at the site known as Proryv Naboko,
were voluminous and highly explosive, with
gas jets erupting >250 m in the air (Gor¬
deev et ak, 2013); later stages produced huge
emissions of sulfur dioxide (SO2), with lava
fountains and pyroclastic deposits. Much
of the early-erupted material consisted of
extremely frothy, glassy lavas and pyroclastic
material. The lavas are silica-rich basalts, and
can be divided into a low-Fe group and a
high-Fe group.
Two samples of the frothy lavas from the
first eruptions yielded abundant diamonds
(>700 grains); most were of a “yellowish-
greenish” colour, and occur in gas cavities
in the lava. Individual grains of moissanite,
corundum (red, blue and Ti-bearing) and
native elements (Fe, Cu, Al) were found in
the same samples, also within gas cavities.
Similar reduced minerals also have been
reported from the lavas of several other vol¬
canos on Kamchatka (Gordeev et ah, 2013),
and both moissanite and Ti-bearing corun¬
dum are common associates of “diamond¬
bearing ores of unconventional types” related
to explosive volcanism in Russia (Karpov et
al., 2014).
27
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
The diamonds from Tolbachik (Karpov et
ah, 2014) are typically 250-700 pm in size,
and show many remarkable similarities with
those from the peridotites of Tibet and the
Polar Urals. They have the same distinctive
colour and the same cubo-octahedral habit,
with smooth cubic faces (Fig. 10); they con¬
tain nitrogen almost entirely as single nitro¬
gen atoms; they have similar isotopically-
light carbon (S^^C = -27 to -23, compared
to -28 to -24 in the Tibetan diamonds).
Some crystals show hollows on the cubic
faces, suggestive of hopper growth (“diffu¬
sion hunger”; Karpov et ah 2014). Unfor¬
tunately, no information is available so far
on the nature of inclusions in the Tolbachik
diamonds.
Figure 10. (a) location of the Kamchatka Peninsula (box) in Far Eastern Russia; (b) map of the
Kamchatka Peninsula showing lines of volcanoes (red dots) related to subduction of the Pacific
Plate beneath the continental margin (Okhotsk Block); red stars mark sites of Avacha volcano,
and the 2012-2014 Tolbachik eruption; (c) SEM images of diamonds from lavas of the Tolbachik
eruption of 2012, showing mixtures of octahedral (111) and cubic faces (cf Fig. 5a); arrows
point to hollow faces that may indicate either hopper growth or later dissolution (After Karpov
et al, 2014).
28
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — - Deep-Earth Methane and Mantle Dynamics
Discussion
Similarities and Differences
(1) Tectonic environment: The three locali¬
ties discussed here lie in three distinct tectonic
environments, but there may be common
factors. The Kamkatcha peninsula lies above
a major ocean-continent collision, with ongo¬
ing subduction and large active volcanoes.
The Tibetan peridotites are in a major conti¬
nental- collision zone (India-Asia), as are the
Polar Urals occurrences (Asia-Europe), and
there is only limited evidence of volcanism
related to their emplacement. However, the
continent-continent collisions in both cases
were preceded by the subduction of oceanic
plates, which may have brought up the peri-
dotite bodies. The Cretaceous volcanism in
the Mt Carmel area also lies along a plate
boundary, but in this case it is the Dead Sea
Transform Fault (Fig. 6), along which the
plates grind past one another rather than sub¬
ducting. In this case the movement on the
fault accommodates the opening of the Red
Sea to the south, transferring the movement
to another major fault in southern Turkey.
This plate boundary (1000 km long) thus
must extend well down into the mantle,
and could provide a conduit for deep-seated
magmas and fluids similar to those generated
by deep subduction.
(2) Associated magmas: The composition
of the lavas that carry the Kamchatka and Mt
Carmel low- jOj mineral assemblages lie in
the broad spectrum of basaltic compositions
associated with shallow melting of the mantle
below the lithosphere. The Kamchatka lavas
are characteristic of the volcanic rocks of
magmatic arcs above continental-margin
subduction zones. The basalts of northern
Israel are more similar to “intraplate” basalts,
and have been described as the products of
a deep-seated mantle plume, with most of
the melting occurring just below the base of
the lithosphere (Fig. Ic; Stein and Hofmann,
1992). The only magmas contemporane¬
ous with the Tibetan peridotites are similar
to the basalts erupted at mid-ocean ridges
(Liu et al, 2015; Zhang et al., 2015); there
is no evidence to link these magmas to the
super-reduced mineral suite. These differ¬
ences suggest that the composition of the
magmas is not a critical factor in generating
the low-^2 assemblages.
(3) Super-reduced mineral assemblages:
The major minerals of the low- j02 assem¬
blages are similar in all three occurrences
(Table 1). The investigations of the Kam¬
chatka lavas have only begun, so relatively
few phases have been identified. The
Tibetan minerals, which have been studied
by many Chinese scientists for more than 30
years, include more than 80 different species,
many known only as single grains (Yang et
al., 2014). The assemblage from northern
Israel has been under detailed study for only
a year or two; it contains >65 identified
compounds, many not previously known
as naturally occurring minerals, but it also
lacks many of those identified in Tibet. This
may reflect the vagaries of observation, but
may also indicate subtle differences between
the conditions of crystallization in different
localities. For example, diamonds are rela¬
tively abundant in the Tibetan and Siberian
localities, but very rare in the Mt. Carmel
volcanics (one microdiamond in a 252 kg
sample); this might reflect differences in the
depth of their respective volcanic systems. It
is important to recognize the unique nature
of the diamonds found in both Tibet/Polar
Urals and Kamchatka, which appears to
require processes different from those that
generate the “normal” kimberlitic diamonds
formed in old continental roots (Howell et
ah, 2015).
29
Journal & Proceedings of the Royal Society of New South ^^^^es
Griffin et aL — Deep-Earth Methane and Mantle Dynamics
Geeeratio.g super-reducing
conditions— some evidenccj
some speculation
As noted in the Introduction, there are two
problems-^ how to generate super-reducing
conditions in the upper mantle, and how
to stop the products from reacting with the
surrounding oxidized mantle, A key to the
first question may lie in the mineral tistarite
(Ti203; Fig. 9). The N. Israel samples rep¬
resent the first known terrestrial occurrence
of this mineral; its “type locality” (Fig. 11)
is a single tiny grain in the Allende mete¬
orite, where it occurs together with TiC
and corundum, as in the N. Israel samples.
Allende is a carbonaceous chondrite— one
of the most primitive types of meteorite, and
one of the oldest objects in the Solar System.
The tistarite-TiC-corundum association in
Allende represents early condensates from
the solar nebula, in regions near the Sun;
they reflect conditions of low pressure, but
high temperature (>1400 °C) and a very low
JO2 (Fig. 2) related to the streaming of the
hydrogen-dominated solar wind through the
nebular cloud. How could we generate such
“nebular” conditions on Earth?
One possibility is illustrated by Figure
2c; in the deep Earth, at ^2 below the IW
buffer, mantle fluids are dominated by meth¬
ane (CH4) and hydrogen (H2). If we can
bring large amounts of such fluids to shallow
depths, and keep them from equilibrating
with the mantle on the way up, they would
have a strongly reducing effect on their
environment when they reached the upper
mantle, as shown by reactions such as (4):
2CH4 + 02^ 2H2 + 2C + 2H2O (4)
where C is diamond or graphite, depending
on pressure.
In addition, reaction (5) can occur at shal¬
low depths (<3 km), and carbon monoxide
Figure 11. (a) Slice of the Allende meteorite,
showing pea-sized chondrules and irregular
light-coloured Calcium-Aluminium Inclu¬
sions, containing refractory phases condensed
from the early Solar nebula; (b) the association
of tistarite (^203), khamrabaevite (TiC) and
corundum in the Allende meteorite (After Ma
and Rossman, 2009). This is the type material
of tistarite, and was the only known occur¬
rence before the discovery of this mineral asso¬
ciation in the corundum from the Mt Carmel
area (c£ Fig. 9).
is a very powerfiil reducing agent in its own
right:
2CH4 + O, 4H2 + 2CO;
2C0->C + H20 (5)
Note that these reactions also can liberate
and deposit large proportions of carbon,
which is consistent with the abundance of
diamonds and/or carbide minerals in these
deposits.
30
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. — Deep-Earth Methane and Mantle Dynamics
Rapid crystallization of corundum, SiC
and other minerals
The second problem noted above is in keep¬
ing the newly-formed low- JO2 minerals
from reacting with the surrounding mantle;
the available observations offer a few sugges¬
tions. Where the Tibetan moissanite (and
diamond) have been seen in situ, the grains
are surrounded by mantles of amorphous
carbon. The nature and origin of this mate¬
rial is not clear from the available research,
but it could at least provide a protective coat¬
ing. Perhaps more important is the presence
in all three localities of a Ti-rich corundum;
our work suggests that this is itself an indica¬
tor of low y02. The hopper growth of the
corundum in the Mt Carmel localities is an
important clue. The rapid growth of a large,
porous network of corundum crystals in a
magmatic (or hydrothermal) conduit could
provide a low- j02 “sheltered environment”
that would be isolated from the surrounding
mantle, while reducing fluids could continue
to flow through the network. This process
could be the key link among the localities.
But what processes can produce a melt
or fluid that is so super-saturated in Al
that it can rapidly crystallize large volumes
of corundum, apparently unaccompanied
by any of the minerals common in mafic
magmas? One possibility is suggested by
the crystallization of anorthite (a feldspar
mineral; CaAl2Si208) in the melt pockets
of the Shefa Yamim corundum. Anorth¬
ite melts directly to liquid, or crystallizes
directly from melts, at low pressure, but at
depths of 10 km-30 km, corundum crys¬
tallizes first from Al-rich melts (Goldsmith,
1980), and anorthite only begins to crys¬
tallize at lower temperatures. Anorthite
is unstable deeper than about 30 km, but
corundum can crystallize from melts to
much greater depths. The simplest way to
make a basaltic magma “dump” corundum
is to remove silica (Si02). If this occurs at
1 0-30 km depths, this desilication can move
the magma into a compositional space rich
in Ca and Al (the anorthite component); at
depths >30 km, corundum will be the first
phase to crystallize, and will continue to be
the only crystallizing phase over a wide span
of temperature as the magma cools. The
reducing processes discussed above provide
a way to desilicate the magma, for example
by reactions like (3) and (6):
Si02 (in silicates) + CH4 SiC + 2 H2O;
H2O + 2CH4 C T CO2 + 4H2 (6)
Thus the streaming of CH4-rich fluids
through the large “plumbing systems” asso¬
ciated with deep-seated volcanism at plate
boundaries could be the process that results
in the crystallization of both corundum
and moissanite, and their associated low-
JO2 minerals, within the upper mantle. In
the case of the Tibetan peridotites, these
plumbing systems have not been recognized;
they may have been obscured by the post¬
emplacement deformation and recrystalli¬
zation of the peridotites and many of the
chromite ores.
Conclusions^ Relationships to
tectonics
It appears that the common factor in the
three localities described here may be the
streaming of CH4 from deep inside Earth’s
mantle, related to deep-seated volcanism
at plate boundaries. In Tibet, geodynamic
models suggest that the peridotites were
exhumed from the Transition Zone (>400
km) at rates of 6™8 cm/year (McGowan et
ah, 2015) — very rapid in geological terms.
We expect that this upwelling process also
would generate large fluxes of deep-man¬
tle fluids, helping to produce the basaltic
magmas. Kamchatka lies above a major
31
Journal & Proceedings of the Royal Society of New South "^^^es
Griffin et aL Deep-Earth Methane and Mantle Dynamics
subduction zone, and the rollback of this
slab would produce similar upwellings from
the deep mantle. In both cases, fluids rich
in CH4 and H2 might also be supplied by
the Mewatering” of carbon-rich marine sedi¬
ments attached to the subducting slab that
triggered the rise of the peridotites.
There was no obvious subduction system
active beneath northern Israel in Cretaceous
time; the magmatism has been ascribed to
the activity of a deep-seated mantle plume,
and it seems likely that such plumes also
would carry deep-mantle, CH4”rich fluids.
However, seismic tomography shows that
the whole Eastern Mediterranean region is
underlain by a network of ancient subducted
slabs, from which buoyant material could
rise, if triggered by tectonic activity or slab
rollback, as in the Tibetan situation. In either
case, local plume-like upwellings could be
focused into the major plate boundary that
became the Dead Sea Transform Fault, pro¬
viding a tectonic environment where both
magmas and deep-mantle fluids could rise
to shallow levels.
We therefore suggest that such ''jets” of
CH4 ±H2 may commonly accompany certain
types of mafic volcanism, especially along
plate boundaries, driven by deep tectonics.
We expect that in the future, the unusual
low- ^2 mineral assemblages described here
will be recognised in many types of volcanic
settings. However, they will only be discov¬
ered when researchers begin to look more
carefully at the minute trace minerals in vol¬
canic ejecta; future studies will reveal if our
speculations are correct.
Acknowledgements
We thank Shefa Yamim Ltd., Dave Apter and
John Ward for their enthusiastic collabora¬
tion on the Mt Carmel material referred to
in this study. Steve Craven gave invaluable
assistance with the mineral separation work.
Shi Rendeng, Qing Xiong and Jonathon
Aitchison are thanked for helpful discus¬
sions on Tibetan geology, and Jingsui Yang
and Paul Robinson for provision of sam¬
ples and published figures relating to the
Tibetan diamond problem. This work was
funded by the ARC Centre of Excellence
for Core to Crust Fluid Systems (CCFS),
with additional support from the Institute
of Tibetan Plateau Research, Chinese Acad¬
emy of Sciences (Beijing). Analytical data
were obtained using instrumentation funded
by DEST Systemic Infrastructure Grants,
ARC LIEF, NCRIS, industry partners and
Macquarie University. This is publication
876 from the ARC Centre of Excellence
for Core to Crust Fluid Systems (http://
www.ccfs.mq.edu.au), and 1126 from the
GEMOC ARC National Key Centre (http://
www.gemoc.mq.edu.au).
References
Afonso, J.C. and Zlotnik, S. 201 1. The
subductibility of continental lithosphere. In
Brown, D. & Ryan, P.D. (Eds), Arc - Continent
Collision. Frontiers in Earth Sciences, Springer-
Verlag, 53-86.
Bai, W-J., Zhou, M-E and Robinson, P-T. 1993.
Possibly diamond-bearing mantle peridotites
and podiform chromites in the Luobusa and
Donqiao ophiolites, Tibet, Canadian Journal
Earth Science 30, 1650-1659.
Dobrzhinetskaya, L.F., Wirth, R., Yang, J-S., Ian,
D., Hutcheon, P.K. and Green, Fl.W, 2009.
High-pressure highly reduced nitrides and
oxides from chromitite of a Tibetan ophiolite.
Proceedings of the National Academy of Sciences
106, 19233-19238.
Garfunkel, Z. 1989. Tectonic setting of
Phanerozoic magmatism in Israel. Israel J.
Earth Science 38, 51-74.
Goldsmith, J.R. 1980. The melting and
breakdown reactions of anorthite at high
pressures and temperatures. American
Mineralogist ■, 272-284.
32
Journal & Proceedings of the Royal Society of New South Wales
Griffin et al. Deep-Earth Methane and Mantle Dynamics
Gordeev, E.L, Muravyov, Y.D., Samoylenko, S.B,,
Volynets, A.O., Melnikov, D.V,, Dvigalo, V.N.
and Melekestsev, LV. 2013. First results from
the 2012-2013 Tolbachik Fissure Eruption Bull
Volcanologkal Society of Japan 58, 1-8.
Griffin, W.L., Afonso, J.C., Belousova, E.A.,
Gain, S.E.M., Gong, X-H., Gonzalez-
Jimenez, J.M., Howell, D., Huang, J-X.,
McGowan, N., Pearson, N.J., Satsukawa, T,
Shi, R., Williams, P., Xiong, Q., Yang, J-S.,
Zhang, M. and O’Reilly, S. Y 2016a. Mantle
Recycling: Transition-Zone metamorphism of
Tibetan ophiolitic peridotites and its tectonic
implications. Journal of Petrology yj ^ 655-684.
Griffin, WL., Gain, S.E.M., Adams, D.T,
Huang, J-X., Saunders, M., Toledo, V., Pearson,
N.J. and O’Reilly, S.Y. 2016b. First terrestrial
occurrence of tistarite (Ti203): Ultra-low
oxygen fugacity in the upper mantle beneath
Mt Carmel, Israel. Geology 44, 8 1 5-8 18.
Howell, D., Griffin, W.L., Gain, S., Stern,
R.A., Huang, J-X., Yang, J., Pearson, N.J. and
O’Reilly, S.Y 2015. Diamonds in ophiolites:
Contamination or a new diamond growth
environment? Earth and Planetary Science
Letters A50, 284-295.
Kaminchik, J. 2014. The origin of intraplate
alkaline mafic magmatism in continental
shelves: lavas and xenoliths from the Upper
Cretaceous volcanoes of Mt Carmel, Israel.
Unpublished MSc thesis, Beer Sheva University,
Israel.
Karpov, G.A., Silev, V.I., Anikin, L.P., Rakin,
V.I., Vasil’ev, E.A., Filatov, S.K., Petrovskii,
V.A. and Flerov, G.B. 2014. Diamonds and
accessory minerals in products of the 20 1 2-
2013 Tolbachik fissure eruption. Journal of
Volcanology and Seismology 8, 323-339.
Liu, E, Yang, J-S., Dilek, Y, Xu, Z-Q., Xu, X-Z.,
Liang, F-H., Chen, S-Y. and Lian, D-Y 2015.
Geochronology and geochemistry of basaltic
lavas in the Dongbo and Purang opniolites
of the Yarlung-Zangbo suture zone: Plume-
influenced continental-margin-type oceanic
lithosphere in southern Tibet. Gondwana
Research 17, 701-718.
Ma, C, and Rossman, G.R. 2009. Tistarite,
Ti203, a new refractory mineral from the
Allende meteorite. American Mineralogist 94,
841-844.
McGowan, N.M., Griffin, W.L., Gonzalez-
Jimenez, J.M., Belousova, E., Afonso., J.C.,
Shi, R., McCammon, C.A., Pearson, N.J. and
O’Reilly, S.Y. 2015. Tibetan chromitites:
excavating the slab graveyard. Geology 43,
179-182.
Mittlefehldt, D.W 1986. Petrology of high
pressure clinopyroxenite series xenoliths. Mount
Carmel, Israel. Contributions to Mineralo^ and
Petrology 94, 245-252
O’Reilly, S.Y. and Griffin, W.L. 2010. Rates
of magma ascent: Constraints from mantle-
derived xenoliths. In Dosseto, A., Turner,
S.P and Van Orman, J.A. (eds) Timescales of
Magmatic Processes: From Core to Atmosphere.
Blackwell Publishing Ltd. Pp. 116-124.
Sass, E. 1980. Late Cretaceous volcanism in
Mount Carmel, Israel. Israel Journal of Earth
Sciences 29, 8-24.
Stein, M., and Hofmann, A.W. 1992. Fossil
plume head beneath the Arabian lithosphere?
Earth and Planetary Science Letters 114, 193-
209.
Xu, X., Yang, J., Robinson, P.T., Xiong, E, Ba,
D. and Guo, G. 2015. Origin of ultrahigh
pressure and highly reduced minerals in
podiform chromitites and associated mantle
peridotites of the Luobusa ophiolite, Tibet.
Gondwana Research 27, 686-700.
Yang, J.S., Dobrzhinetskaya, L., Bai, W-J., Fang,
Q-S., Robinson, P.T, Zhang, J. and Green,
H.W II 2007. Diamond- and coesite-bearing
chromitites from the Luobusa ophiolite, Tibet.
Geology y, 875-878.
Yang, J.S., Robinson, P.T. and Dilek, Y. 2014.
Diamonds in ophiolites. Elements 10 127-130.
Yang, J.S., Meng,. E, Xu, S., Robinson,
P.T, Dilek, Y, Makeyev, A.B., Wirth, R.,
Wiedenbeck, M. and Cliff, J. 2015. Diamonds,
native elements and metal alloys from
chromitites of the Ray-Iz ophiolite of the Polar
Urals. Gondwana Research 27, 459-485.
Zhang, C., Liu, C-Z., Wu, F-Y, Zhang, L-L.
and Ji, W-Q. 2015. Geochemistry and
geochronology of mafic rocks from the Luobusa
Ophiolite, South Tibet. Lithos 245, 93-108.
33
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 34-50. ISSN 0035-9173/16/010034-17
Chemoenzymatic pathways for the synthesis of biologically
active natural products
Martin G. Banwell*, Benoit Bolte, Joshua N. Buckler, Ee Ling Chang,
Ping Lan, Ehab S. Taher, Lorenzo V. White and Anthony C. Willis
Research School of Chemistry, Institute of Advanced Studies,
Tlie Australian National University, Canberra, ACT 260 1 , Australia
* Corresponding author.
Email: Martin.Banwell@anu.edu.au
.Abstract
The whole-cell biotransformation of mono-nuclear aromatic compounds using certain genetically-
engineered micro-organisms that over-express the enzyme toluene dioxygenase (TDO) allows for the
large scale production of compounds known as r/j-l,2-dihydrocatechols. These metabolites, which
are normally obtained in enantiomerically pure form, can be manipulated, by chemical means, in a
range of distinct (and predictable) ways with the result that they have proven to be especially versatile
starting materials for the assembly of a range of structurally diverse and biologically active systems.
Herein we describe, on a case-by-case basis, the recent applications of various combinations of
TDO-mediated and chemical steps in so-called chemoenzymatic total syntheses of a range of organic
compounds with therapeutic potential
Introduction
hemical space the space spanned by
all possible s,mall molecules and chemi¬
cal compounds) is essentially infinite.^ The
challenge, then, has been to access the most
meaningful or useful parts of it. Nature
has provided critical inspirations. So, 3.8
billion years of evolution has produced a
global molecular library of unsurpassed size,
structural diversity and functional value —
our planet’s chemome.23,4 Humankind has
sought to "'mine” this bioactive molecule
resource for its benefit and such endeav¬
ors have been spectacularly successful as
evidenced by the existence of the remark¬
able array of medicines, materials and agro¬
chemicals that underpin society as we know
it today. As a result the world we live in has
been transformed. This is evidenced by our
exploitation of drugs with household names
such as penicillin, morphine and Taxol®.
There are many additional but perhaps less
well-known examples. For instance, organ
transplant surgery would fail completely
without the post-operative application of
the chemome-derived anti-rejection drugs
such FK506 and cyclosporin A.^ Similarly,
a significant number of agents that control
agricultural pests, and so helping to ensure
botli tlie security and efficiency of world
food production, have also come from
Nature/ the global chemome.^
Despite such successes, enormous chal¬
lenges remain. So-called unmet scientific
and societal needs include those arising
from the development of resistance to cur¬
rent therapies (perhaps seen most promi¬
nently in the area of antibiotics^) and, in the
34
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al— Chemoenzymatic Pathways
agrochemical sector, pest-control agents.^ In
addition, there is a desperate need for small
molecule entities that provide, inter alia^
effective control of neurodegenerative
diseases and diabetes in a globally aging
population, for ones that treat certain types of
refractory cancers and for others that
effectively modulate mammalian and other
immune systems.
After forays into areas such as
combinatorial chemistry,^ major players in the
pharmaceutical industry, sometimes in
parmerships with Government- funded
agencies, are remrning to interrogation of the
chemome (or at least portions thereof) as a
means for productively probing chemical and
thence biological space. There are a number
of reasons for such moves^® including the
recognition that, for example, the current
pharmaceutical industry is built on <10% of
the biosynthetic capacity of the microbial
world, one that continues to show a
remarkable ability to deliver biologically
relevant small molecules.
Occurring in tandem with these trends is
the emergence of a plethora of new
techniques and concepts concerned with the
generation of biologically relevant molecular
diversity involving the use of, inter alia^
techniques of de novo biosynthesis for
producing functionally annotated chemome
components, creation of new metabolic
pathways,i2 synthetic fermentation, and
activity-directed synthesis.i^ Simultaneously,
new synergies are being recognized between
in vitro, in vivo and in silico studies of drug
metabolism and thus allowing for much
more efficient/ rapid assessments of the utility
of certain compounds as molecular probes,
drugs and/ or agrochemicals.
The development of new methods and
protocols for effecting the chemical synthesis
of biologically active natural products and
various analogues remain important parts of
the range of activities concerned with
exploiting components of the global
chemome for therapeutic and other purposes.
At least two motivations drive such efforts,
the first being the need to address issues of
supply. Thus, it is often the case that
secondary metabolites^ are only available in
miniscule amounts from their natural source
with the result that insufficient material is
available for development purposes.
Chemical synthesis is often the best method
for addressing such issues. Secondly, tmly
useful chemical syntheses offer the capacity to
generate analogues of the natural product that
would not normally be available through
manipulation of the natural product itself
This article, which is based on a lecture
presented by the senior author at the
University of Sydney as part of the RSNSW’s
2014 Liversidge Award, details work being
undertaken at the Australian National
University on the exploitation of certain
chemoenzymatic methods for the synthesis of
biologically active natural products and thek
analogues. The work is presented according
to the class of namral product being targeted
as well as the stmctural and chemical
relationships between them.
Results and discussion
The term chemoenzymatic synthesis used in
this article, and elsewhere, refers to the
assembly of target compounds using a
combination of chemical and enzymatic
techniques. While there are many variations
on this theme that reflect the extraordinarily
diverse range of chemical and enzymatic
transformations available these days, the
specific form of the latter that appHes here
involves the whole-ceU bio transformation of
a range of simple and readily available
aromatic compounds of the general form 1
(Scheme 1) into the corresponding <^>-1,2-
dihydrocatechols (2).^^ When genetically
engineered micro-organisms such as E. coli
JM109 (pDTG601)i2 ^re used for such
35
jouRN.'iL & Proceedings of the Royal Society of New South Wales
Banwell et al— Chemoenzymatic Pathways
purposes, these metabolites can be readily
produced at kilogram scales and are often
obtained in >99.95 enantiomeric excess (ee).
In the illustrated cases the enzyme
responsible for these conversions is toluene
dioxygenase (TDO) but a number of related
ones are known including biphenyl
dioxygenase, naphthalene dioxygenase and
toluate dioxygenase. The end result is that a
remarkable suite of ahl ,2-dihydrocatechols
and related metabolites is known - these
number in the many hundreds at the present
time.^^*^ Given the capacities to produce
numerous mutants, and thus expand the
range of substrates that can be
biotrans formed, the possible extensions of
such processes would appear to be vast. A
further fascinating aspect of them is the
“chemoselectivities” they can display. So, for
example, styrene (1, X = CH=CH2) is
converted into the triene 2 (X = CH=CH2), a
process wherein the aromatic ring is oxidised
in preference to the exocyclic olefin, a
fonctional group selectivity that cannot be
achieved by any of the strictly chemical
methods known at the present time.^^
Pseudomonas
putida
39»D
or
E. COUJM109
(pDTG601)
X
1
2
>99.8% ee
X = H, Me, Cl, Br, I, CN, CH=CH2, CF3, COjH etc
Scheme 1
The utility of the aYl,2"dihydrocatechols (2)
as starting materials in chemical synthesis has
taken some time to be recognised in a
broader sense. Various groups, especially
those led by Ley in the and Hudlicky in
North Americad^^’^ have carried out the
pioneering work in the area. Such studies
established the reactivity “patterns” shown in
Figure 1 as well as attendant hazards arising
from the dehydrative re-aromatisation of
these substrates^o and the propensity of
certain derivatives, most notably the
corresponding acetonides, to engage in
normally unproductive Diels-Alder (DA)
dimerization reactionsdi
jmorGin’dereZhydroxyl)
Hazard:
dehydration/aromatisation
more nucleophilic
double-bond
(when X = halogen)
allylic alcohols
H
Claisen-type
rearrangments
Figure 1
Our own contributions in tiie area began in
the late 1 9808^2 and in the intervening period
we have been able to establish a series of total
syntheses (Figure 2) that emphasise the
extraordinary range of natural product targets
available through manipulation of these
metabolites. Some specific examples arising
from our recent research are discussed on a
case^by^case basis in the following sections.
36
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al.- Chemoenzymatic Pathways
NHAc
^ ^NH2*H3P04
lycorine
degradation product
'tt.
H02C.„_
(-)-tricholomenyn A
OH O
HO2C— d />“OH
H .0
HO HN
(+)-brunsvigme
OMe MeO
HOv.A..„OH
OH
(+)-armillarivm
platencin
(-)-phomentrioloxin
Figure 2
The Ribisins
Ribisins A-D were isolated by Fukuyama and
co-workers from Phellmus ribis (Schmach.)
Qu£ (Hymenochaetaceae),“3 a hingus used in
traditional medicine for various purposes.
Using a range of spectroscopic methods they
were assigned structures 3-6 (Figure 3),
respectively, and shown to enhance neurite
outgrowth in PC 12 cells at ca. 1 pM
concentrations. As such they have potential
for development as agents for the treatment
of certain neurological disorders.
37
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al™ Chemoenzymatic Pathways
Given the structural resemblance of the
polyoxygenated cyclohexane ring of these
natural products to the
dihydrocatechols 2 (X = Br) we sought a
means for effecting the relevant chemical
conversions. The route used for establishing
a synthesis of compound 5, the structure
assigned to ribisin C and the most active
compound in the series, is shown in Scheme
2.24
x-Q
t)H
HO
2 (X = Br)
(i) 2,2nMP,
l?-TsOH
(ii) ot-CPBAs
DCM
■'-(7
aq. HCl,
THF
(90% over two steps)
pH
OH
X
(i) CH3I, NaH,
THF
(ii) AcOH/HjO
,-p-
hA
,OMe
j OMe
OH
(81% over three steps)
OH
111
14 (49% over two steps)
Swem
oxidation
Scheme 2
38
Journal & Proceedings of the Royal Society of New South W7\les
Banwell et al.— Chemoenzymatic Pathways
The opening stages of this reaction sequence
are typical of the manner in which the as-1,2-
dihydrocatechols can be manipulated and
involve the initial conversion of compound 2
(X = Br) into the corresponding acetonide
and the regio- and stereo-selective
epoxidation of the latter to give the oxirane 7.
Treatment of compound 7 with aqueous
mineral acid resulted in a regioselective ring¬
opening reaction to afford the trans-^o\ 8 that
could be bis-O-methylated under conven¬
tional conditions and the resulting acetonide
was then cleaved, again under conventional
conditions, to give the <7>-diol 9 that embodies
most of the key elements of the Eastern
hemisphere of target 3. Compound 9 could
be engaged in a Suzuki-Miyaura cross¬
coupling reaction with the commercially
available boronate ester 10 and two products
thereby formed, namely the bis-phenol 11 and
the dihydrobenzohiran 12. Product 12 is
presumably formed through cycHsation of the
initially produced cross-coupHng product
while congener 11 arises from successive loss
of the elements of water and methanol (no
particular order implied) from the same
intermediate. The lone hydroxyl group within
compound 11 could be protected as the
corresponding a-chloro-acetate 13, a
necessary step because of the looming
introduction of a second hydroxyl group as
the precursor to the ketone moiety. The use
of the a-chloroacetate as a protecting group
proved essential as in the final step of the
reaction sequence attempts to remove the less
labde parent acetate resulted in
decomposition of the substrate. Epoxidation
of compound 13 using ;5^-chloroperben2oic
acid (;^-CPBA) led, presumably via
spontaneous rearrangement of the initially
formed oxirane, to the benzofiiran alcohol 14
that could be oxidised to the corresponding
ketone 15 under Swern conditions. Cleavage
of the a-chloroacetate residue within this last
compound was accomplished using zinc
acetate in methanol and thus forming the
target compound 5. While aU the usual
spectroscopic data acquired on compound 5
matched those reported for ribisin C, the
specific rotation derived from the synthetic
material was of the same magnitude but the
opposite sign to that reported for the natural
product. The implications are clear — the
structure of ribisin C is represented by
stmcture entS rather than 5. Since we
required an authentic sample of ribisin C {ent-
5) for biological testing, a synthesis of it was
pursued. This could be achieved (Scheme 3)
using the same starting material and many of
the same transformations as employed in
generating its enantiomer (5). A key step of
the fourteen-stage reaction sequence involved
the inversion of configuration at C3 within a
derivative of compound 2 (X = Br) using
IVIitsunobu chemistry. As a result ribisin C
was obtained and all of the derived data,
including the specific rotation, matched those
reported for the natural product.
fourteen
steps
Mitsunobu reaction
using CICH2CO2H as
nucleophile
ent-S
[I2]d = - 10.8(c 0.5,MeOH)
Lit. value [0]d = - 1 1.1 (c 0.5, MeOH)
Scheme 3
Extensions of this sort of chemistry enabled
the synthesis of all of the stmctures originally
assigned to the ribisins and thus revealed that
while ribisins A and D are constituted as
originally described-^ that attributed to
congener B is, Hke C, incorrect-s The true
stmctures of all the ribisins are shown in
Figure 4 with the corrected stereocentres
within compounds B (16) and C {ent-S)
highlighted in red. Extensive biological
evaluations of the ribisins and the range of
39
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al — Chemoenzymatic Pathways
congeners prepared during the course of our
synthetic studies are now underway.
ribisin A (3)
ribisin C (ent-S)
ribisin B (16)
ribisin D (6)
Figure 4
Analogues of Galanthamine
Ribisin D (6) bears a ‘‘provocative” structural
resemblance to the ABC ring-system of the
alkaloid galanthamine (17) that is used in
many countries for the symptomatic
treatment of Alzheimer’s disease (Figure
As such we were prompted to explore means
by which the chemistry described above
could be adapted so as to produce
compounds bearing greater similarities to
galanthamine (or, in the first instance at least,
the enantiomer thereof.
ribisin D (6) 17
Figure 5
An efficient reaction sequence leading to a
dioxygenated derivative of ^///-galanthamine is
shown in Scheme 4“"^ and involves an initial
reaction of the abovementioned oxitane 7
with ^-methoxybenzyl alcohol (^-MBOH) in
the presence of boron trifluoride diethyl
etherate to generate the anticipated addition
product that upon treatment, in a second
step, with methanol containing pyridinium p~
toluenesulfonate (PPTS) affords triol 18. This
last compound could be converted into the
corresponding Ley ketal 19-^ through
treatment with 2,2,3, 3-tetramethoxybutane
(2,2,3,3-TMB) in the presence of p-
toluenesulfonic acid ^-TsOH) /trimethyl
orthoformate (TMOF) and Suzuki-Miyaura
cross-coupHng of this with the boronate ester
20 (produced dicectly from 0-methoxyphenol
using a C-H functionalization protocol)
afforded the arylated cyclohexene 21. This last
compound that was itself engaged in an
intramolecular Mitsunobu reaction using di-
iso-propjl azodicarboxylate (DLAD) to afford
the dihydrobenzofuran 22.
Despite tlie potential for aromatisation
(through simple double-bond migration),
compound 22 could be engaged in an
Eschenmoser-variant of the Claisen
rearrangement reaction using the dimethyl
acetal of ATdV-dimethylacetamide-^ and thus
affording the angularly substituted ABC-ring
analogue 23 of ^///-galanthamine. Over three
conventional steps compound 23 could be
converted into its mono-methylated
counterpart 24. The last compound
participated in a Pictet-Spengler cycHsation
reaction on exposure to a mixture of
paraformaldehyde and trifluoroacetic acid
(TEA) and the presumably first-formed
product 25 underwent cleavage of the Ley
acetal residue to give diol 26 as the only
isolable product of reaction. Compound 26,
representing a dioxygenated derivative of ent-
galanthamine {ent-YT)^ and various congeners
that have been prepared using related reaction
sequences are currently being subjected to
evaluation as inhibitors of the neurologically
significant enzyme acetylcholine esterase
(AChE).
40
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al™ Chemoenzymatic Pathways
(i) PMBOH,
OPMB
BF.
OH
(ii) PPTS
MeOH
HO OH
18 (70%)
2,2,3,3-TMB
i?-TsOH, TMOF
MeOH
HO O
OMe
MeO
19 (86%)
OMe
MeO OH HO O
MeO
Ph3P, DIAD,
21 (71%)
Pd(dppf)Cl2*DCM
EtjN
THF/HjO
THF
OH
(Ty / \ ,xOMe
MeC(OMe)2NMe2
O
MejN--^^
/r**\ /
1 ''
MeO
-
toluene
yOi-
MeO
reflux
MeO M
22 (96%)
23 (86%)
(CHO)„
TFA, DCE
Me O
three steps|
O
Scheme 4
26 (63%)
It is worth noting, at this point, that the
enantiomer of certain of the
dihydrocatechols described above are also
available.^o So, for example, biotransform¬
ation of ^-iodotoluene or ^-iodobromo-
benzene [27a (X = Me) and 27b (X = Br),
respectively] (Scheme 5) using E. coli]MlQ9
(pDTG601) affords metabolite 28 that upon
exposure to dihydrogen in the presence of
palladium on carbon undergoes
hydrogenolytic cleavage of the associated C-I
bond and thus delivering either dsA,2-
dihydrocatechol ent-2 (X = Me) or ent-2 (X
Br).
27
Scheme 5
E. coliJM109
(PDTG601J
X = Me or Br
2
(original metabolite)
ent-2
41
jouRN.\L & Proceedings of the Royal Society of New South Wales
Banwell et aL— Chemoenzymatic Pathways
The Opiates
Vinblastine, Vincristine and VindoMne
Morphine and its congener codeine are
members of opiate family. They are used
extensively for the management of pain and
represent the most widely applied and highest
grossing medicines in the world today.^^ Their
structural complexity means that for the
moment, at least, opiates such as morphine
are obtained from natural sources and then
derivatized by simple chemical means so as to
produce related drugs. Nevertheless, much
progress has been made in terms of
developing commercially viable total
syntheses of these systems. HudMcky and co^-
workers have defined the current “gold
standard” in the area.^^ Given the tantalising
structural resemblance between the readily
available compound 23 and ^»Acodeine (29)
(Figure 6) we are now attempting to modify
the synthesis of the former so as to access the
latter. This will likely involve introducing the
necessary additional two-carbon unit by using
a variant of boronate ester 20 and completing
the synthesis of the less fiinctionalised
cyclohexane ring within target 29 using an
intramolecular Sn^ reaction that
simultaneously cleaves the Ley acetal subunit.
OMe
Vinblastine (30) and vincristine (31) (Figure
7) are indole-indoline-based alkaloids derived
from various plant sources, perhaps most
notably the Madagascan rosy periwinkle.^^
They are used in the clinical treatment of
non-Hodgkin's lymphomas as well as
testicular, breast and lung cancers. These
compounds are derived in vivo from the
significandy more abundant and co-occurring
alkaloid vindoline (32), Given the
development of direct, chemically based and
“bio-inspired” methods for effecting the
conversion of this simpler compound into
alkaloids 30 and 31, vindoline has become the
focus of considerable attention as a synthetic
target,34
Our own efforts in this area have been
inspired by the observation (Figure 8)^^ ^p^t
the mutant organism P. putida BGXMl can
effect, in an enantioselective fashion, the
whole-cell bio transformation of abundant m-
ethyltoluene (33) into the carboxylic add diol
34 that bears a striking resemblance to the
highly fijnctionaMsed C-ring of vindoline.
Accordingly, a recent focus of some of our
work in the area of chemoenzymatic synthesis
has been on identifying methods for
converting this metabolite into vindoline (32)
and thence into vinblastine (30) and
vincristine (31).
23 29
fe«f-codeine)
Figure 6
Madagascan rosy periwinkle
42
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al— Chemoenzymatic Pathways
Figure 7
Figure 8
The model study outlined in Scheme 6 has
provided encouragement.^^ Thus, the ^Yl,2-
dihydrocatechol 2 (X = Br), representing a
model for congener 34, was converted, by the
means described earlier, into the oxirane 7.
Treatment of this last compound with acetic
acid in the presence of mineral acid afforded a
trans-dkA mono-ester that was protected
under standard conditions as the
corresponding MOM-ether and thus
affording compound 35 that could be cross-
coupled with ethyl boronic acid in the
presence of a Pd[0] catalyst to give, after
completing cleavage of the acetate residue
using methanolic potassium carbonate, the
allylic alcohol 36. This last compound was
engaged in a sluggish Eschenmoser-Claisen
rearrangement reaction to give amide 37, the
side-chain of which could be elaborated, over
three steps, into the nitrile 38. Over a further
three conventional steps this was converted
into the a-iodocyclohexenone 39 that itself
served as a substrate for a palladium-catalysed
UUmann cross-coupling reaction^^ with o-
iodonitroarene 40 and so delivering the a-
arylated cyclohexenone 41. On exposure to
dihydrogen in the presence of Raney cobalti^
and a proton source compound 41 engaged in
a series of chemoselective reductions and two
cyclisation reactions with the result that the
tetracycHc compound 42 was formed. The
completion of the synthesis of the pentacyclic
framework of vindoline proved straight¬
forward and involved reaction of the last
compound with 2-bromoethanol in the
presence of base, mesylation of the resulting
alcohol 43 and treatment of the sulfonate
ester so formed with potassium /(?/f-butoxide
to generate the isoindole 44.
In an effort to introduce the
carbomethoxy group associated with alkaloid
32, compound 44 was subjected to successive
treatment with /(?//-butyllithium then Mander's
reagent (NCC02Me).3^ However, rather than
obtaining the hoped-for C-carbomethoxy-
lated imine, carbamate 45 was produced,
presumably by a pathway whereby the tert-
butyllithium acts as a hydride source^o with
the resulting indoHne anion then reacting (at
nitrogen) with the added electrophile. Efforts
are now underway to adapt these chemistries
so as to convert metabolite 34 into vindoline.
The most challenging issue associated with
doing so will be finding a means for
introducing the C-C double bond
incorporated within the D-ring of target 32.
43
Journal & Proceedings of the Royal Society of New South Wales
Banwell et aL— Chemoenzymatic Pathways
2 (X = Br)
(i) 2,2-DMP,
l?-TsOH
(ii) m-CPBA,
DCM
(i) AcOH/H'"
(ii) MOM^Cl/base
7 (90% over two steps)
Br
OMOM
3S (89%)
(i) EtB(OH)2, I 00 K2CO3,
Pd[0] j MeOH
NC
OMOM
38 (71%)
three steps
MeC(OMe)2NMe2
heat
Raney
aey cobalt | MeOH
MeO
(OMsCI.EtjN jjQ-
(ii) r-BuOK
MeO
44 (73%)
43 (92%)
t-BuLi
then NCCOjMe
Scheme 6
The ProtoiHudanes
The title sesquiterpenes embody a distinctive
tricyclic framework wherein a central
cyclohexane ring is annulated, in an angular
arrangement, to both a four- and a five-
membered ring.^^ The protoilludane aryl
ester (+)-armillarivin (46) (Figure 9) has been
found in the edible sugar mushroom
A.rmiUaria melk^- while representative
additional natural products in this family
include 4743 ^nd 48^4 that are derived from
the saprotrophic wood decomposing fungus
Granukhasidium velkreum (ElHs & Cragin)
JuDlick
44
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al.-- Chemoenzymatic Pathways
Figure 9
In 2013 we described the first and thus far
only total synthesis of (+)-armiUarivin.45 A key
element of our approach (Scheme 7) was an
initial high-pressure promoted and completely
regio- and stereo-selective Diels-Alder
reaction between the as-1 ,2-dihydrocatechol 2
(X = Me) and cyclopentenone.^^ Relatively
conventional but rather extensive
manipulations of the resulting adduct lead to
the cyclop entannulated bicyclo [2.2.2] octen-
one 49 that engaged, as a second pivotal step
of the synthesis, in a photochemicaUy-
promoted 1,3-acyl migration reaction (Givens
rearrangement) to afford the tricyclic isomer
50. This last compound, which embodies the
tricyclic protoiUudane framework, was readily
manipulated over just three steps to deliver
(+)-armillarivin. The structure of this
synthetically produced material was
confirmed by single-crystal X-ray analysis and
aU the derived spectroscopic data, including
specific rotation, matched those reported for
the natural product.
Subjection of the acetonide derivative of
compound 2 (X = Me) to a Diels-Alder
reaction with cyclopentenone affords, vm
addition of the dienophile to the face of the
diene opposite to that “occupied” by the
hydroxyl groups, cyclopentannulated bicyclo-
[2.2.2] octenones that are enantiomericaUy
related to those obtained by the pathway
described immediately above. In essence,
dien, by controlling the facial selectivity of
such cycloaddition reactions eidier
enantiomeric form of the relevant Diels-Alder
adduct can be obtained.
sixteen steps
including intital
Diels-Alder reaction
2 (X = Me)
reductionyesterification
Scheme 7
45
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al.— Chemoenzymatic Pathways
By such means we have recently been able to
complete total syntheses of the enantiomeric
forms of the protoilludanes 47 and 48^8 and
thus confirmingj for the first time, the
structures assigned to them.
Platencin
The Diels-Alder cycloaddition chemistry
involving aTl,2"dihydrocatechols as the
component can be effectively extended to
intramolecular variants. This is perhaps best
exemplified in our recently completed firsU
and second“generation chemoenzymatic
syntiieses of platendn (51)549,50 ^ compound
isolated from Streptomyces platensis MA7327
that acts as a potent and dual inhibitor of
FabH and FabF, key enzymes associated with
fatty acid biosynthesis in bacteria (Figure
10) By virtue of its novel structure and
modes of action, platencin is regarded as an
important new lead in the development of
urgently needed, nexUgeneration anti=
bacterial agents4“
In our first generation synthesis of
compound 51 (Scheme 8)5^9 the acetonide
derivative, 52, of the aTl,2-dihydrocatechol 2
(X = I) was engaged in a Stille cross-coupling
reaction with the Z-configured
aU^enylstannane 53 to give tlie tetra-ene 54.
Substrate 53 was prepared in a
straightforward manner with the
stereochemistry at the quaternary carbon
centre being controlled through the agency of
a chiral auxiliary.
WWle compound 54 failed to engage in
an intramolecular Diels-Alder (IMDA)
reaction, the readily derived ketone 55 did so
when heated in refluxing toluene and thus
affording, in stereochemically pure form,
adduct 56 embodying the tricarbocyclic core
of platendn. Over a further thirteen steps
compound 56 could be converted into (“■)-
platendn (51).
Some of these steps were needed to deal
with functional group incompatibilities, an
issue that has been addressed, albeit in a
modest way, through our recently disclosed
second-generation synthesis.^® In a devel¬
oping collaboration with the Hudlicky group
at Brock University (Canada), efforts are now
focussed on a third-generation approach.
Figure 10
46
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al— Chemoenzymatic Pathways
51 56 (70%)
[(-)-platencin]
Scheme 8
Conclusions
Enzymes have an almost unparalleled
capacity to transform simple organic
substrates into synthetically more valuable
ones, especially enantiomericaUy pure
compounds (EPCs). Through the use of
various genetic engineering, gene shuffling
and directed evolution techniques the
opportunities to expand upon the existing
‘Tbrary” of metabolites seem almost infinite.
Furthermore, pathway-engineering techniques
provide a capacity to produce (mutant)
organisms that effect, through the
orchestrated action of a series of enzymes,
multistep transformations. The conversion
of /W-ethyltoluene (33) into compound 34^^ is
a case in point and wherein both mono¬
oxygenases and dioxygenases act in concert to
produce a potentially high-value metaboHte.
When combined with the power of chemical
synthesis (as manifest in the techniques of
chemoenzymatic synthesis), such bio¬
transformations provide a powerful tool kit
for preparing a wide range of compounds of
biological relevance. Ironically, perhaps,
while microbiologists have a remarkable
capacity to generate a diversity of low
molecular weight metabolites (and often at
multi-kilogram or larger scale) and synthesis
chemists have an almost insatiable appetite
for new synthons, the often siloed nature of
academic research activities results in less than
desirable overlap of the relevant sets of
expertise. Changing this situation can only
benefit both discipHnes.
Acknowledgements
We thank the Australian Research Council
and the Institute of Advanced Studies at the
AustraKan National University for ongoing
support. The contributions of our colleagues
who co-authored the publications referenced
below are also gratefully acknowledged, as are
the useful comments of Rob Capon and
Craig Williams (University of Queensland),
Ron Quinn (Griffith University) and Peter
Kamso (Macquarie University).
47
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al — Chemoenzymatic Pathways
References and notes
1 J. L. Reymond, Chem. R^s., 2015, 48, 722„
2. The term chemome is used here in the sense
defined by Wender (see re£ 3) and taken to
mean that vast collection of secondary
metabolites produced by the planet’s flora and
fauna.
3. P. A. Wender, Nal Prod 2014, 31, 433.
4. The temi “terpenome” has been introduced to
define that subset of the chemome that is
terpenoid in origin: M. B. Quin, C. M. Flynn and
C. Schmidt-Dannert, Nat. Prod. 2014, 31,
1449.
5. See, for example, H. van Hattum and H.
Waldmann, Am. Chem. Soc., 2014, 136, 11853;
(b) S. Rizzo and H. Waldmann, Chem. R£p., 2014,
114, 4621; (c) M. E. Maier, 0^. BiomoL Chem.,
2015, 13, 5302.
6. B. C. Gerwick and T. C. Sparks, Pest Manag. Set.,
2014, 70,1169.
7. M. A. T. Blaskovich, J. Zuegg, A. G, Elliot and
M. A. Coo^&T, ACS Infect. DA, 2015, 1, 285.
8. R. F. Service, Science, 2013, 341, 1329.
9. A very limited number of drugs has thus far
emerged from the de novo applications of
combinatorial chemistry techniques: D. J.
Newman and G. M. Cra^, J. Nat. Prod., 2012,
75,311.
10. (a) E. Kellenberger, A. Hofmann and R. J.
Quinn, Nat. Prod. Rfp., 2011, 28, 1483; (b) J. L.
Fox, Nat. Biotech., 2014, 32, 305; (c) A. L. Harvey,
R. Edrada-Ebel and R. J. Quinn, Nat Rev. Drug
Discov., 2015, 14, 111; (d) M. Pascolutti, M.
CampitelJi, B. Nguyen, N. Pham, A.-D. Gorse
and R. J. Quinn, PPmS ONE, 2015, 10,
e0120942.
11. J. R. Doroghazi, J. C. Albright, A. W. Goering,
K.=S. Ju, R. R. Haines, K. A. Tchalukov, D. P.
Labeda, N. L. KeUeher and W. W. Metcalf,
Nature Chem. Biol, 2014, 10, 963.
12. x. Zhu, J. Liu and W. Zhang, Nat Chem. Biol,
2015, 11,115.
13. Y. Huang and J. W. Bode, Nat Chem., 2014, 6,
877.
14. G. Karageorgis, S. Warriner and A. Nelson,
Nat Chem., 2014, 6, 872.
15J. Kirchmair, A. H. GoUer, D. Lang, J. Kunze,
B. Testa, 1. D. Wilson, R. C. Glen and G.
Schneider, Nat Rev. Drug Discov., 2015, 14, 387.
16. For reviews on methods for generating cis~\,2-
dihydrocatechols by microbial dihydroxylation
of the corresponding aromatics, as well as the
synthetic applications of these metabolites, see:
(a) T. Hudlicky, D. Gonzalez and D. T. Gibson,
AMrichimica Acta, 1999, 32, 35; (b) M. G.
Banwell, A. J. Edwards, G. J. Harfoot, K. A.
Jollffe, M. D. McLeod, K. J. McRae, S. G.
Stewart and M. Voegtie, Pure AppL Chem., 2003,
75, 223; (c) R. A. Johnson, Org. React, 2004, 63,
117; (d) T. Hudlicky and J. W. Reed, Sjn/ett,
2009, 685; (e) D. J.-Y. D. Bon, B. Lee, M. G.
Banwell and L A. Cade, Chimka O^i, 2012, 30,
No. 5, (Chiral Technologies Supplement), 22; {£)
U. Rinner, Chiral Pool Synthesis: Chiral Pool
Syntheses from aYCyclohexadiene Diols In: E.
M. Carreira and H. Yamamoto, (Eds)
Comprehensive Chirality, 2012, 2, 240.
17. G. J. Zylstra and D. T. Gibson,/. Biol Chem.,
1989, 264,14940.
18. The chemical dihydroxylation of benzene has
been reported as a means of preparing the
cyclitol (±)-pimtol: P. M. J. Jung, W. B.
Motlierwell and A. S. Williams, Chem. Commun.,
1997, 1283.
19. See, for example, S. V. Ley, F. Sternfeld and S.
Taylor, Tetrahedron Rett., 1987, 28, 225.
20. (a) S. M. Brown and T. Hudlicky, in Organic
Synthesis: Theory and Applications, 1992, JAI
Press Inc., Greenwich, Connecticut, 1992, voL
II, 113; (b) D. R. Boyd, J. Blacker, B. Bryne, H.
Dalton, M. V. Hand, S. C. Kelly, R. A. More
O’Ferral, S. N. Rao, N. D, Sharma and G. N.
Sheldrake, /. Chem. Soc., Chem. Commun., 1994,
313.
21. These types of acetonides are prone to
dimerization: (a) S. V. Ley, A. J. Redgrave, S. C.
Taylor, S. Ahmed and D. W. Ribbons, Sjnktt,
1991, 741; (b) T. HudHcky, E. E. Boros, H. F.
Olivo and J. S. Merola, / Org. Chem., 1992, 57,
1026.
22. M. G. Banwell, Org. Prp. Proced. Int, 1989, 21,
255.
23. Y. Liu, M. Kubo and Y. Fukuyama, / Nat
Prod, 2012, 75,2152.
24. P. Lan, M. G. BanweU, J. S. Ward and A. C.
Willis, Org. Lett., 2014, 16, 228.
25. P. Lan, M. G. Banwell and A. C. Willis, J. Ofg.
Chem., 2014, 79, 2829.
26. For points-of-entiy into the relevant literature
48
Journal & Proceedings of the Royal Society of New South Wales
Banwell et al.— Chemoenzymatic Pathways
see: (a) J. Nugent, E. Matousova and M. G.
Banwell, Eur. J. Org, Chem.^ 2015, 3771; (b) M. G.
Banwell,}. Buckler, C. J. Jackson, P. Lan, X. Ma,
E. Matousova and J. Nugent J. — Devising New
Syntheses of the Alkaloid Galanthamine, a
Potent and Clinically Deployed Inhibitor of
Acetylcholine Esterase in Strategies and Tactics in
Organic Synthesis (Ed. M. Harmarta), 2015 11, 29.
27. J. N. Buckler, E. S. Taher and M. G. BanweU,
unpublished observations.
28. S. V. Ley, D. K. BaeschHn, D. J. Dixon, A. C.
Foster, S. J. Ince, H. W. M. Priepke and D. J.
Reynolds, Chem. Rer., 2001, 101, 53.
29. For a relevant application of this process see
M. G. Banwell, X. Ma, O. P. Karunaratne and A.
C. Willis, /. Chenf., 2010, 63, 1437.
30. (a) D. R. Boyd, N. D. Sharma, S. A. Barr, H.
Dalton, J. Chima, G. Whited, R. Seemayer, J.
Am. Chem. Soc., 1994, 116, 1147; (b) C. C. R.
Allen, D. R. Boyd, H. Dalton, N. D. Sharma, 1.
Brannigan, N. A. Kerley, G. N. Sheldrake, S. C.
Taylor,/. Chem. Soc., Chem. Commun.^ 1995, 117.
31. For relevant background accounts see: (a) L.
D. Kapoor, Opium Poppy: Botany, Chemisty, and
Phannacokgg, Food Products Press, New York,
1997; (b) M. Booth, Opium: A Histoy, St.
Martin’s Press, New York, 1998.
32. J. W. Reed and T. Hudlicky, Acc. Chem. Res.,
2015, 48, 674 and references cited therein.
33. (a) M. E. Kuehne and 1. Marko, The Alkaloids,
1990, 37, 77; (b) H. L. Pearce, The Alkaloids,
1990; 37, 145; (c) N. Neuss and M. N. Neuss,
The Alkaloids, 1990, 37, 229.
34. J. E. Sears and D. L. Boger, Acc. Chem. Res.,
2015, 48, 653.
35. M. G. Banwell, A. J. Edwards, D. W. Lupton
and G. Whited, J. Chem., 2005, 58, 14.
36. L. V. White and M. G. Banwell,/. Org. Chem.,
2016, 81,1617.
37. M. G. BanweE, M. T. Jones and T. A. Reekie,
Chem. N. Z., 2011, 75, 122.
38. M. G. BanweU, M. T. Jones, T. A. Reekie, B.
D. Schwartz, S. H. Tan and L. V. White, Ofg.
Biomol Chem., 2014, 12, 7433.
39. S. R. Crabtree, W. L. A. Chu and L. N.
Mander, Synlett, 1990, 169.
40. G. B. Bennett, W. J. HouHhan and R. B.
Mason,/ Organomet. Chem., 1975, 99, 185.
41. For a recent and comprehensive review of the
area, see P. Siengalewicz, J. Mulzer and U.
Rinner, Eur. J. Ofg. Chem., 2011, 7041.
42. (a) J. S. Yang, Y. L. Su, Y. L. Wang, X. Z.
Feng, D. Q. Yu, and X. T. Liang, Yaoxue Xuebao
(Acta Phamaceutica Sinica), 1991, 26, 117; (b) P.
Cremin, D. M. X. DonneUy, J.-L. Wolfender and
K. Hostettmann, / Chromatogr. A, 1995, 710,
273; (c) D. M. X. DonneUy, T. Konishi, O.
Dunne and P. Cremin, Phytochem., 1997, 44,
1473.
43. C. L. Nord, A. Menkis, R. Vasaitis and A.
Broberg, Phytoche?n., 2013, 90, 128.
44. C. L. Nord, A. Menkis, C. Lendel, R. Vasaitis
and A. Broberg, Phytochem., 2014, 102, 197.
45. B. D. Schwartz, E. Matousova, R. White, M.
G. BanweU and A. C. WiUis, Org. Lett., 2013, 15,
1934.
46. See, for example, M. G. BanweU, K. A. B.
Austin and A. C. WUUs, Tetrahedron, 2007, 63,
6388.
47. For a review of this and related
photochemicaUy-promoted rearrangements see
M. G. BanweU and D. J.-Y. D Bon, AppUcations
of the Di-rt-Methane and Related
Rearrangement Reactions in Chemical Synthesis
in Molecular Rfarrangements in Ofganic Synthesis (Ed.
C. M. Rojas), 2015, 261.
48. E. L. Chang, B. Bolte, P. Lan, A. C. WiUis and
M. G. BanweE, J. Org. Chem., 2016, 81, 2078.
49. E. L. Chang, B. D. Schwartz, A. G. Draffan,
M. G. BanweU and A. C. WiUis, Chem. Asian /.,
2015, 10, 427.
50. R. N. Muhammad, A. G. Draffan, M. G.
BanweU, M. G. and A. C. WUUs, Synlett, 2016, 27,
61.
51. For useful reviews of this topic see (a) K.
Tiefenbacher and J. Mulzer, Angew. Chem. Int
Ed, 2008, 47, 2548; (b) K. Palanichamy and K.
P. KaUappan, Chem. Asian /., 2010, 5, 668; (c) E.
Martens and A. L. Deamin,/ Antibiot., 2011, 64,
705; (d) M. Saleem, H. Hussain, 1. Ahmed, T.
van Ree, and K. Krohn, Nat. Prod. Rep., 2011, 28,
1534; (e) A. M. AUahverdiyev, M. Bagirova, E. S.
Abamor, S. C. Ates, R. C. Koc, M. Miralogu, S.
Elcicek, S. Yaman and G. Unal, Infect. Drug.
Resist. 2013, 6, 99.
52. (a) R. E. W. Hancock, Nat. Rsv. Drug Discov.,
2007, 6, 28; (b) P. C. Appelbaum,/ Antimicrob.
Chemother., 2012, 67, 2062; (c) S. Shapiro,/.
Antibiot, 2013, 66, 371; (d) R. Tommasi, D. G.
49
JouRN.^L & Proceedings of the Roy.4l Society of New South Wales
Banwell et al — Chemoenzymatic Pathways
Brown, G. K. Walkup, J. 1. Manchester and A.
A. Miller, Nat. Rep. DmgDiscov.., 2015, 14, 529.
Martin Banwell is a Professor of Chemistry in
the Research School of Chemistry at the
Australian National University. His research
focus is on the total synthesis of biolo^cally
active natural products and his contributions
in this area have formed the basis of his
award of the 2014 Liversidge Lectureship and
Medal.
50
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 51-58. ISSN 0035-9173/16/010051-08
The curious economist: William Stanley Jevons in
Sydney
Address at the Dinner to Mark the Opening of the Exhibition at the
Powerhouse Museum^
The late Ian Casdes^
Former Australian Statistician, 28 October 2004
Abstract
In 2004, which marked the 150th anniversary of William Stanley Jevons arrival in Sydney,
the Powerhouse Museum mounted an exhibition on “Jevons: The Curious Economist/’ Jevons
was born in Liverpool, in England. He studied chemistry in London, and came to New South
Wales at the age of 1 9 to work at the Sydney Mint. The Gold Rush was in full swing, and his
job was to assess the quality and purity of the precious metal from the diggings. This work put
him in a unique position, at the intersection of geology, chemistry, economics and industry.
Wanting to document as many aspects of life in Australia as he could, Jevons also took up
photography, even taking his camera equipment to the goldfields. After five years in Sydney,
Jevons returned to England, where he eventually became the first Professor of Economics at
University College in London. He became a member of the N.S.W. Philosophical Society on
13 June 1856. He left Australia in April 1859. He drowned in 1882, aged 46.
At a dinner to celebrate the opening of the Exhibition, the late Ian Castles gave this address.
Ian Castles (1935-2010) was an accomplished statistician and civil servant. He was Secretary
of the Commonwealth Department of Finance (1979-1986), the Australian Statistician (1986-
1994), and Visiting Fellow at A.N.U. between 1995 and 2000. He was Executive Director
and Vice President of the Academy of the Social Sciences in Australia. His family has given
permission for the Journal to publish the address, lightly edited by Robert Marks.
Introduction
begin by congratulating the Powerhouse
Museum and especially the exhibition
curators — Matthew Connell and Lindsay
^ http://www.powerhousemuseum.com/ exhibitions/
jevons. php
2 Lightly edited, with references, by Robert Marks,
2016.
Barrett — on the splendid exhibition we’ve
had the privilege of viewing this evening.
I’m sure that the curators would agree that
they had in ‘'the curious economist” an
extraordinary subject upon which to focus.
My job tonight is to try and give some
added context to the amazing story that
began just 150 years ago when the teen¬
age Stanley Jevons landed in this city. Em
51
Journal & Proceedings of the Royal Society of New South Wales
Castles — William Stanley Jevons in Sydney
honoured that the Powerhouse has invited
me to perform this task, and especially
grateful in diat, ten or twelve years ago, the
Museum could only have seen me as a nagger
and a nuisance.
My interest in persuading the
Powerhouse to mount an exhibition on
Jevons’ years in Sydney dates back to a
conference that I attended at the Reserve
Bank in the early 1990s. I can’t remember the
subject of the conference, but I remember
well that, during the luncheon break, I strolled
southward along Macquarie Street with a
visiting British economist. I pointed across
the street to the fine building that formerly
housed the Sydney branch of the Royal Mint,
and remarked tiiat that was where Jevons had
spent the first four or five years of his
working Hfe.
My companion’s initial reaction was one
of disbelief So we crossed the road and spent
a few minutes in the building known as the
Mint Museum, which at that time was part of
the Powerhouse Museum. To my satisfaction,
we were able to confirm immediately that
Jevons had indeed worked at the Mint.
But to my consternation, and that of my
economist colleague, the captions to the
display revealed nothing about who Jevons
was or what he had done. Let me quote:
This room ... focuses on the lives of two
Mint employees and social aspects of the
gold msh era. [It] focuses on contrasting
experiences of those years: life on the
diggings and the experiences of two
gentlemen who worked at the Mint,
William Jevons and Robert Hunt...
This was tme as far as it went, but my
companion and I thought that visitors to the
Mint Museum should have been told that
‘"Wniiam Jevons” was later to become one of
the founders of modern economics. And
perhaps also that he pursued a remarkable
range of interests in his Sydney years.
So strongly did I believe this that I wrote
to the director of the Powerhouse Museum to
suggest that the a museum of applied arts and
sciences was missing an opportunity by not
making a good deal more of the young
Jevons’ association with Sydney — and his
contribution to the applied arts and sciences
during his five years in this city. I had a polite
response, and one or two useful meetings
with people at the Powerhouse.
But, just at this stage, the Powerhouse
Museum announced a major acquisition: the
famous difference engine invented by Charles
Babbage, commonly recognised as the
foremnner to the modern computer. In
discussions at the Powerhouse, I said that this
was all very well, and that the engine was
indeed an eminently appropriate artefact for a
museum of applied arts and sciences. But
Babbage had had no connection with
Australia.
I argued that the Museum should take an
interest in the fact that Jevons too had built a
proto-computer. Moreover, unlike Babbage,
the former resident of Sydney had done the
whole thing at his own expense. And Jevons’
machine was the first machine that could
solve a complex logical problem faster than
that problem could be solved without using
the machine.
Then there was a further development.
The Powerhouse Museum acquired Jevons’
telescope — not just a telescope like the one
Jevons used, but the instmment that he
actually used. I was told, and of course I
agreed, that one object directly relating to
Jevons was not enough to build an exhibition
around. The inscription on the telescope
reveals that, not long after leaving Sydney,
Jevons gave it to his cousin who was then in
Penang in what is now Malaysia. It is not clear
how or why this telescope found its way back
to Australia.
52
Journal & Proceedings of the Royal Society of New South Wales
Castles — William Stanley Jevons in Sydney
The instmment is, I suggest, a deeply
symbolic artefact. It encapsulates Jevons’
passionate interest in the means and meaning
of observation and measurement of the
natural and the social world. It was through
this telescope that the young Jevons
whom we now know to have been Australia’s
first urban sociologist ■ — first viewed an
Australian city. The city was Melbourne, not
Sydney. In a ship anchored off the port of
Melbourne on 24 September 1854, Jevons
confided to his journal that:
the town is very curious; it looks like a
crowd of ugly buildings of every size
chiefly iron and wood arranged as if no
two houses were in one street. There
seem to be very few large buildings and
these are as ugly as the rest, but we can
make out by the telescope an Exhibition
building, 2 or 3 churches ... [and] a
railway station on the other side of the
bay. (Black & Konekamp 1972, p. 109)
This would have been the jusucompleted
collection of weatherboard train sheds on the
site now occupied by Flinders Street station.
In Sydney, Jevons used the same Httle
telescope to observe two eclipses of the sun
in 1857. In an extended report written after
the second of these events, he told the
readers of Henry Parkes’ newspaper. The
Empire, that
The first contact of the moon with the
lower limb of the sun could be easily
observed with the aid of an ordinary
telescope, and seemed to take place
within a few seconds after the time ...
calculated by Mr. Tebbutt, of Windsor,
my watch being regulated by the noon
gun at Fort Philip. The coincidence
would, no doubt, have been the more
complete the more accurate the means of
observation. (The Empire, 19 Sept. 1857,
p.4)
“Mr. TebbutT, of Windsor” was later to
become the world-famous amateur
astronomer who formerly appeared on the
Australian $100 banknote. At this time, he
was 23 years of age, one year older than
Jevons. The report of the eclipse in The
Empire on the day after the eclipse is quite an
extended one, and Jevons’ diary on the day
that he wrote this report survives in the John
Rylands library in Manchester.
Let me quote the entry for this day in the
life of W. S. Jevons, a 22-year-old public
servant:
After sleepless night got up about 3.30
and started to Bellevue HiU in dark.
About 5 a.m. commenced observations
concerning eclipse. Returned to Mint
[this was presumably where he wrote the
report that was published on the
following day]. Tea at Mr. Newton’s
fMr.Newton was Chief Engineer at the
Mint] and then to Victoria Theatre.
Brooke and Airs. Hek very good in
Much Ado About Nothing.
Nature books for WiQy Newton 8/6
[presumably his host’s son], WheweU’s
Lect. on Political Economy 11/-.
On the facing page of the diary, the entry
for the succeeding day (28 March 1857) reads:
Played much music in evening [this
would have been on the harmonium that
appears in the photograph of his study at
Double Bay, which appears in the
exhibition]. Meteorological work, etc.
Wrote and sent letter to Empke with aim
^ Like Jevons, John Tebbutt Jr. was also a member of
the Philosophical Society of N.S.W.
53
jouRN.-\L & Proceedings of the Royal Society of New South Wales
Castles ~ William Stanley Jevons in Sydney
of shutting up writers about “Protective
humbug”^.
Jevons was a strong free-trader.
During Jevons’ stay in Sydney, The
Empire published many reports and letters
authored by him, on subjects as diverse as
meteorology; the land and railway policy of
New South Wales; the resolutions passed at
the protection meeting; “A Cure for the
Revenue”; lead poison in the Sydney water;
the danger from lightning strikes on the
ammunition stores on Goat Island; a new
sun-gauge^; the royal prerogative of mercy;
the conflict between the archaelogical and
Biblical accounts of the length of human
existence on earth; and “New Facts
Concerning the Interior of AustraHa’’.^"
Many of Jevons’ pieces in The Empire
were quite combative. He acknowledged this
in a letter to his cousin toward the end of his
sojourn in Sydney (9 October 1858):
I am becoming quite accustomed to the
pen as a weapon of offence & defence,
indeed I suffer under such a rage for
writing that I am scarcely to be tmsted
with a pen in my hand. I often write a
newspaper article and am then on thorns
for ever so long after for fear of a Hbel
action. But alas! my organ the ’Empire’
has passed away in bankruptcy and I am
now reduced to that milk & water affair
the “Herald” which too has not yet
learned to appreciate me, and sticks to
small type, on the backpages, whereas
Parkes always gave me large type next
after the leading article, & usually
reprinted me for the monthly summary.
4 See his letter to The Empire of March 30, 1 857, listed
below.
5 Jevons' paper, “On a sun-gauge, or new actinometer,”
was presented at the Philosophical Society of N.S.W. on
8 July 1857.
^ See the Reference list below for many of these, with
links.
I send you the monthly summary of the
Herald which is certainly a wondertiil
publication for a single day; the number
of copies sold too is something
enormous. You wiU find in it every scrap
of colonial news... (Black 1973-1981, 2,
pp. 345-346).
Some of these comments are somewhat
unfair to the Herald, which had done Jevons
proud by publishing his long and fascinating
review of the first consolidated volume of the
monthly Sydney Magazine of Science and Art,
to which Jevons himself had made significant
contributions.’^
In the event, it has turned out to be for
the best that the plans for an exhibition on
Jevons in Sydney, including especially the
Symposium, have come to fruition in 2004
rather than in 1994. Because, in the
intervening ten years, Jevons’ life and work in
Sydney and afterwards has attracted
unprecedented attention from researchers in a
wide range of disciplines, both in Australia
and overseas. At least a dozen peer-reviewed
journals have published papers focusing
solely or mainly on aspects of Jevons’ work.
The kst of the names of the journals in which
these smdies have appeared testifies to the
diversity of Jevons’ interests, and of the
continuing scholarly interest in Jevons:®
The EListory of Political Economy,
The History of Economics Eeview,
The Journal of the Histoy of Economic
Thought,
European Journal of the Histoy of Economic
Thought,
Economic Inquiy,
'' Sjdn^ MomingHerald, 27 Sept. 1858, p.8.
8 Ian Castles did not specify which papers the following
list refers to, but in the References below I have included
possible papers — Ed.
54
Journal & Proceedings of the Royal Society of New South Wales
Castles ~ William Stanley Jevons in Sydney
Measurement in Phjsics and Economics
Discussion Papers at the London School of
Economics,
The Histog and Philosophy of Logic,
A.ustralasian Music Pesearch,
The Australian Meteorological Maga^ne,
The Journal of Physical Oceanography,
and
Scientific American.
I would like to be able to talk about the
significance of some of this work, much of
which has been produced by one or other of
the distinguished scholars who are with us
this evening and will be presenting papers at
tomorrow’s symposium.^ But it would be
beyond my capacity to do these scholars
justice, and I can only urge those who may be
able to come to the symposium to do so.
The primary source material for those
who are engaged in research on Jevons
consists, of course, of his own writings,
published and unpublished. Only a small
fraction of his writings in Australia appeared
in print during his lifetime. In a volume
published soon after his death, Jevons’ widow
published edited extracts from his personal
journal and from many of the surviving letters
(Jevons 1886). Much more extensive extracts
of his papers and correspondence have been
published in subsequent years.
Yet much of Jevons’ Australian work
remains unpublished. In a paper published as
recently as 1999, the leading Jevons scholar
Bert Mosseknans and a Belgian colleague
(Mosselmans & Mathijs 1999) published
extracts from a manuscript by Jevons —
uncompleted but mnning to 50 pages —
entitled “On the Science and Art of Music”
^ Mike White, Monash; Harro Maas, University of
Amsterdam; Lindsay Barrett, University of Western
Sydney; Geoff Barker, University of Sydney; Megan
Martin, Historic Houses Trust NSW; Neville NichoUs,
Bureau of Meteorology; Matthew ConneU, Powerhouse
Museum; Jamie Kassler, Australian Academy of
Humanities; Ian Castles, A. N. U.
which is located at the John Rylands Library
at Manchester University. The work was
written in Sydney and sent to Jevons’ sister in
London as each chapter was completed. The
John Rylands Library^o ^nd the Public Library
of New South Wales hold many letters and
other manuscripts relating to Jevons’
Australian years which have never been
published.
It is a pity that much of Jevons’ writings
in Australia — whether published in
contemporary journals and newspapers,
intended for publication in Sydney or in
Britain, or never intended for publication at
aU ■ — is not now readily available. They are of
value not only as a record of several formative
years in Jevons’ own development, but to
tiiose interested in the history of Sydney and
of Australia at this time.
It is now possible for manuscript
material to be “digitised”, enabling it to be
disseminated freely. This has the further
advantage, by comparison with publication in
hard copy, of providing the user access to
interactive facilities for selective searching and
retrieval of the contents.
So I am led to conclude with a
suggestion. I’ll introduce it by pointing out
that, some thirty years ago, the Royal
Economic Society felt justified in publishing
hundreds of pages of letters and papers
written by Jevons while in Australia, in the
handsome seven-volume set edited by
Professor Collison Black. It is fait to say that
much of this material is of interest to
Australian social historians but sheds little
light upon the history of economic thought:
this is tme, for example, of the accounts of
Jevons’ journeys to the Hunter Valley and
Maitland, the Macquarie towns , the
http:/ /www.library^manchester.ac.uk/ search-
resources/ guide-to-special-coUections/ atoz/jevons-
family-papers/
Black & Konekamp 1972, pp. 123-130, 23 May 1856.
55
JouRN.\L & Proceedings of the Royal Society of New South Wales
Castles ~ William Stanley Jevons in Sydney
lUawarra district Bathurst and Sofala^^^
Braidwood and Araluen and the Victorian
gold fields^ Ironically, this wealth of material
is Utde known in Australia except among
those who have been led to it by their interest
in the life and work, especially the economic
work, of Stanley Jevons.
With the advances of technology that
have now occurred, the opportunity now
arises to link all of the Australian-related
Jevons material that is readily available in
published sources with the as-yet unpublished
letters, papers and photographs that are held
in research libraries. The result would be an
interactive product of considerable potential
interest and value, and not only to scientists
of all colours. The cost would not be small,
but nor need it necessarily be prohibitive. If
the exhibition that opens tomorrow
stimulates as much interest in ""the curious
economist” in Sydney as I hope that it wiU, I
believe that the publication of a consolidated
archive in digitised form of the surviving
record of Jevons’ years in Sydney, published
and unpublished, would be a worthy and
imaginative project. It is something that the
Powerhouse Museum and its supporters
might wish to consider.
I am confident that such a project would
attract international interest — and not only
among research scholars but among people in
many walks of Hfe who find fascination in the
development of an extraordinary mind
one of the minds of the century, as John
Maynard Keynes wrote to Lytton Strachey in
1905, when Keynes was 22.
12 Black & Konekamp 1972, pp. 1344 54, 24-28
December 1856.
13 Black & Konekamp 1972, pp. 159-178, 9-13 April
1856.
14 Black & Konekamp 1972, pp. 213-238, 9 March
1856.
13 Three letters to his sister, Lucy Jevons, 13 and 16
March 1859 and 9 April 1859, reprinted in Black
1973-1981,2, pp. 366-373.
In the Preface to her book M World 'Ruled
bj Number, the first fuU-length study of the life
and work of William Stanley Jevons, Dr.
Margaret Schabas, now Professor of
Philosophy at the University of British
Columbia, wrote that she ""had found, over
the course of six or seven years, that my views
on these subjects have changed considerably
and that Jevons had an even richer set of
insights than I had initially suspected... That
the world is ruled by number may still be
subject to dispute, but not the element of
simplicity and beauty in Jevons’s vision of
mind and matter.”
Thank you again for inUting me to speak
this evening. I wish the Exhibition and the
accompanying Symposium every success.
Primary Writings
Black, R. D. C. and Kdnekamp R. (ed.), (1972).
Papers and Correspondence of William Stankj Jevons.
Volume 1. Biographj and Personal Journal London:
Macmillan in association with the Royal
Economic Society.
Black, R. D. C. (ed.), (1973-1981). Papers and
Correspondence of William Stankj Jevons. Volumes
20 7. London: Macmillan in association with the
Royal Economic Society.
Jevons, H. A. (ed.), (1886). Letters and Journal ofW.
Stankj Jevons. London: Macmillan.
Jevons, W. S. Protection, [a letter], The Empire,
Sydney, 30 March, 1857, p.3
http:/ / trove.nla.gov.au/ newspaper/ article/ 6027
3487
Jevons, W. S. Comparison of the land and railway
policy of New South Wales, The Empire, Sydney,
8 April 1857, p.6
http:/ / trove.nla.gov.au/ newspaper/ article/ 6027
_ 6432
Jevons, W. S. On the resolutions passed at the
Protection Meeting, [a letter]. The Empire,
Sydney, 4 May 1857, p.3.
http:/ / trove.nla.gov.au/ newspaper/ article/ 6027
4810
Jevons, W. S. The public lands of New South
Wales. The Empire, Sydney, 24 June 1857, p.4
56
Journal & Proceedings of the Royal Society of New South Wales
Castles - William Stanley Jevons in Sydney
http:/ / trove.nla.gov.au/ newspaper/ article/ 6498
4880
Jevons, W. S. On a sun-gauge, or new actinometen
The Empire, Sydney, 14 July 1857, p.3
http:/ / trove.nla.gov.au/ newspaper/ article/ 6498
5980
Jevons, W. S. The eclipse. The Empire, Sydney, 19
September 1857, p.4.
http:/ / trove.nla.gov.au/ newspaper/ article/ 6026
3467
Jevons, W. S. Lead poison in the Sydney water, [a
letter]. The Empire, Sydney, 5 October 1857, p.5.
http:/ / trove.nla.gov.au/ newspaper/ article/ 6026
6124
Jevons, W. S. Gunpowder and lightning, [a letter].
The Empire, Sydney, 14 November 1857, p.3.
http:/ / trove.nla.gov.au/ newspaper/ article/ 6026
1514
Jevons, W. S. A cure for the Revenue. The Empire,
Sydney, 14 April 1858, p.4.
http:/ / trove.nla.gov.au/ newspaper/ article/ 6042
7881
Jevons, W. S. The royal prerogative of mercy. The
E?7jpire, Sydney, 10 June 1858, p.3.
http:/ / trove.nla.gov.au/ newspaper/ article/ 6042
5807
Jevons, W. S. New facts concerning the interior of
Australia. The Empire, Sydney, 10 August 1858,
p.3.
http:/ / trove.nla.gov.au/ newspaper/ article/ 6042
4807
Jevons, W. S. Review of The Sydney Maga^ne of
Science and Art. Vol 1. Sydney: James W. Waugh,
1858. The Sydney Morining Herald, 27 September
1858, p.8.
http:/ / trove.nla.gov.au/ newspaper/ article/ 1 302
0816
Day & Day 1984 record that Jevons was admitted
as a member of the Philosophical Society of
N.S.W. at its second meeting, 13 June 1856.^*^
He remained a member until he left the colony
for England, in March, 1859.
See also The Empire of 14 June 1856, p.3.
http:/ / trove.nla.gov.au/ newspaper/ article/ 60249242
References.
Bowman, Rhead S. (1997) Policy implications of
W. S. Jevons’s economic theory. Journal of the
Histoy of Economic Thought, 19, 02, 196-221. Day,
A. A. and Day, J. A. F. (1984). A biographical
register of members of the AustraKan
Philosophical Society (1850L5J and the
Philosophical Society of New South Wales
(1856-66). Part 1. Journal and Proceedings of the Royal
Society of New South Wales, 117: 119-127. http:/ /
biodiversipEbraiT.org/ page/ 46349357
Flatau, Paul (2004) Jevon’s one great disciple:
Wicksteed and the Jevonian revolution in the
second generation. History of Economics Review, 40:
69-107, Summer.
Kassler, Jamie C. (1996) William Stanley Jevons:
music and the mechanisation of logic.
Australasian Music Research, 1 .
Maas, Harro (1999) Mechanical rationality: Jevons
and the making of economic man, Studies in
History and Philosophy of Science, Part A 30 (4):
587-619.
McChesney, Fred S., Shughart, William F. II, and
Haddock, David D. (2004) On the internal
contradictions of the law of one price. Economic
Inquiry, 42.4: 706-716.
Mosseknans, Bert (1998) William Stanley Jevons
and the extent of meaning in logic and
economics. History and Philosophy ofEogic, 19, 2,
83-99, January.
Mosseknans, Bert and Mathijs, Ernest (1999)
Jevon’s music manuscript and the political
economy of music, Histoiy of Political Economy,
suppL Economic Engagements with Art, 31: 121-156.
NichoUs, N (1998) William Stanley Jevons and the
climate of Australia, Australian Meteorological
Magacfne, 47, pp. 285-93.
Owen, David (2010) The efficiency dilemma. The
New Yorker, Dec. 20 & 27.
http: / / www.newyorker.com/ m2Lg2am.tl 2010/1
2/ 20/ the-efficiency-dilemma
Schabas, Margaret (1990) A World Railed ly Number:
William Stanley Jevons and the Rise of Mathematical
Economics, Princeton University Press.
Schabas, Margaret (1999) The economics of W. S.
Jevons, History of Political Economy, 31.3: 604-605.
Schmitt, Raymond W. (1995) The salt finger
experiments of Jevons (1857) and Rayleigh
(1880), The Journal of Physical Oceanography, 25:
8-1 7, January 1995.
57
JouRN.\L & Proceedings of the Royal Society of New South Wales
Castles — William Stanley Jevons in Sydney
Steedman, Ian (1997) Jevon’s Theory of Political
Economy and the ’Marginalist Revolution,’
'European Journal of the History of Economic Thought,
4, 1, 43-65, Spring.
White, Michael V. (2006) Cultivated circles of the
Empire: Bibliographical notes on W.S. Jevons’s
antipodean interlude (1854-1859), Histoy of
Economics Review, AT. 101-122.
58
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 59-64. ISSN 0035-9173/16/010059-06
William Stanley Jevons,
Fellow of the Philosophical Society of New South Wales,
1856-1859
Robert E. Marks
Economics, University ofN.S.W.
Email: robert.marks@gmaiLcom
Abstract
This paper attempts to accomplish three things: to report on Jevons’ activities while in Sydney, specifi¬
cally with the Philosophical Society; to argue that his activities in Sydney led dkectly to his work in
economic theory and application on his return to England; and to underhne Jevons’ achievements in
independently pioneering what is now known as “neo-classical” microeconomics, along with writers
in France and Austria. It complements Castles’ 2004 address (Castles, 2016).
Introduction
illiam Stanley Jevons v^as born in Liv¬
erpool on September 1, 1835. His
father, Thomas Jevons, was an iron merchant
and engineer who constructed one of the
first iron boats. His maternal grandfather
was the historian William Roscoe. Aged
16, he entered University College, London,
and studied chemistry and botany In 1853,
before graduating, he accepted a well-paid
post as assayer at the new Sydney Mint on
Macquarie Street, possibly as a consequence
of his fathers business having collapsed. He
arrived in Melbourne on September 24,
1854, and in Sydney soon after.
He threw himself into the intellectual
Hfe of the colony: he collected and studied
local plants; he documented the geology of
the Hawkesbury basin; he wrote the first
study of Australasian weather patterns; he
researched the formation of clouds; he
made pioneering “social surveys” of Sydney
and Goulburn; he debated the economics of
railway construction and land allocation in
the pages of the newspapers. ^
It is recorded that Jevons attended
meetings of the Philosophical Society on
the following dates: 13 June 1856, 8 July
1857, and 9 December 1857, and later in
the monthly Sydney Magayine of Science and
A.rts\ photographs of his were included in a
Photographic Exhibition of the Philosophi¬
cal Society at the Australian Library on 19
December 1859, although he had left Aus¬
tralia months previously, returning to Eng¬
land via North America.
While in Sydney Jevons kept a detailed
journal and diaries, and sent regular letters
to his cousin, his sisters, his brother, and
his father, describing the environment, his
activities, his ideas, and innermost thoughts.
These have since been published, first, by
his widow, in 1886, and, then, in several vol¬
umes, by the Royal Economic Society, in
1973-81.
1 He was not above sending the odd riddle to the
papers: “Why ought the officers of the Mint to be
prosecuted for treason? Because they are constantly
engaged in carrying out a design upon the sovereign.
— J. Jevons, Royal Mint.” Sydney Morning Herald, 20
Jan. 1859, p. 5.
59
Journal & Proceedings of the Royal Society of New South Wales
Marks — William Stanley Jevons FPhSN 1856-59
Consequently, we have a reasonable idea of
the development of his intellectual life during
his Sydney sojourn.
So what influence did his time in Sydney
have on Jevons’ thinking, and what role did it
play in bringing liis ideas to fruition? In fact,
Jevons said later that nearly aU his ideas came
to him during the period he spent in Sydney.
The new colonial city certainly gave him the
time, the money and the intellectual freedom
to undertake work in a wide range of areas:
meteorology, photography, geography,
geology, botany and natural science and a
social survey of the city itself
From Black (1973), we know that on
June 25, 1854, he writes, though I have
got a pretty complete set of common
chemical Apparatus and chemicals, I feel as if
I should take more to Geology & botany,
though I don’t at all intend to stick to Science
above everything else all my life” (Black, p.
78). On October 31, 1855, he talks of
intending to compile and send
“Meteorological Reports to some paper”
(Black, p. 198); the first such published report
appeared in the Empire on 3 September 1856.
On July 19, 1856, the Sydney Morning
Herald published his first letter on “the
science of economy,” about the need for
profitability in railway construction (Black,
pp. 235). This was in reaction to the
Governor’s declaration “that a railway need
not necessarily be capable of paying any
profits, since indirect benefits to the
population may repay its costs.” (Black, p.
265). Sir WilHam Denison had been an officer
in the Royal Engineers.
On October 21, 1856, Jevons writes that
“a Scientific education is one of the best
things possible ... It tends to give your
opinions and thoughts a sort of certainty ^ force ^
and clearness which forms an excellent
foundation for other sorts of knowledge less
precisely determined and established” (Black,
p. 244). Apart from his meteorological work,
he was very interested in botany, music, and
the theatre.
On April 4, 1857, Jevons writes “I have
been much occupied lately with a new Subject
viz. Political Economy, which seems to
mostly suit my exact method of thought.”
(Black, p. 280). He had read Smith’s Wealth of
Nations, as well as more recent books, and had
just published a letter in the Empire “on some
questions of Economy here” (against
protection) and followed that with a letter on
the need for “wise regulations on the sale or
distribution of unoccupied lands of the
colony, and of a right poHcy in the formation
of railways” (Black, pp. 282). On June 17, he
writes “the subject I have been most of all
concerned in for the last sik months is
Political Economy,” having read books by
Smith, Chambers, Martineau, Mill, Whately,
and Chaning. (Black, p. 292)
In a letter of February 28, 1858, Jevons
expounds to his sister: '‘Economy, scientifically
speaking, is a very contracted science; it is in
fact a sort of vague mathematics which
calculates the causes and effects of man’s
industry, and shows how it may best be
applied. ... I have an idea ... that my insight
into the foundations and nature of the
knowledge of man is deeper than that of
most men or writers ... To extend and perfect
the abstract or the detailed and practical
knowledge of man and society is perhaps the
most useful and necessary work in which any
one can now engage. There are plenty of
people engaged with physical science, and
practical science and arts may be left to look
after themselves, but thoroughly to
understand the principles of society appears
to me now to be the most cogent business.”
(Black, pp. 321).
On June 9, 1 858, he says: “Social science
is the wide subject before me, and I have
2 See also the Extracts from the Personal Diaries, R. D.
C Black (ed), G?/ KTJ, 1981, pp. 115.
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Marks — William Stanley Jevons FPhSN 1856-59
even had for many years the idea of a work
on 'Towns & Cities,’ to analyse their
constitution, and causes, the relative character
of their parts, & the relative character of
particular cities & thus eventually lead to such
knowledge of their nature & shall ensure their
improvement, as any Scientific knowledge is
eventually reduced to practice.” (Black, pp.
327).
On August 4, 1858, Jevons says, 'Tt
seems to me that Man is a subject as little
understood now as the Heavens (Astronomy)
were by the Ancients. Within the last hundred
years, sciences almost innumerable have
spmng up, but mostly devoted to physical
Nature. Comparatively few have perceived
that Human Namre may also be the subject
of a science. It is indeed a many-sided subject.
Religion, metaphysics, ethics, jurispmdence,
political economy, politics, & even, medicine,
art, poetry and many other studies aU have
man for the subject. But the social condition
of man as influenced by the many internal &
external circumstances is perhaps an
indefinite but a wide & rich field for future
research.” (Black, pp. 335).
In his penultimate letter from Sydney, on
January 30, 1859, Jevons asks his sister, "...
does it not strike you that just as in Physical
Science there are general & profound
principles deducible from a great number of
physical phenomena, so in treating of Man or
Society there must also be general principles
and laws which underlie all the present
discussions & partial arguments? ... Man is
said to possess free will but however this may
be, he is at least a phenomenon in which
effect is always connected with cause. All the
investigations of Social Science must proceed
on the assumption that there are causes as
make people good & bad, happy & miserable,
rich & poor, as weU as strong & feeble. It
foUows that each individual man must be a
creature of cause (& effect. ... To attempt to
define the foundations of our knowledge of
man, is surely a work worth a lifetime ...”
He socialised with members of the
Philosophical Society: on 13 July 1857 he
spent the day "taking a bush walk with the
old Rev. Mr. Clarke, the geologist, afterwards
dining with him, & Mrs. Clarke, and the two
Miss Clarkes.” (Black, p. 298). It was,
ironically, W. B. Clarke, who, as president of
the Society, urged it in a direction away from
Jevons’ emerging interests: for the next
century it focussed very much on the physical
sciences, largely excluding economics and
other social sciences.
So he returned to England, completed
his B.A. and M.A. degrees, and was elected to
a chair as the professor of logic and mental
and moral philosophy and Cobden professor
of political economy at Owens College,
Manchester, in 1866. Meanwhile, in Australia
he was remembered as the inventor of a sun
gauge (1857), and the author of a contentious
study of clouds (1857) and of The
Climatology of Australasia (1859), as well as
being a pioneering photographer (Bourke,
1955). Later, the newspapers reported his
analysis of gold prices (1863), and his
influential book on the effects of the
exhaustion of British coal mines (1866).
Australian newspapers were, however, almost
mute on his advances in theoretical
economics.
On the basis of entries in Jevons’ diary
for 1860, La Nauxe (1953) states that "young
Jevons arrived on one identifiable day,
Febmary 19, 1860, at a comprehension of the
true Theory of TcommyT This is less than twelve
months after his leaving Sydney. In a letter to
his brother, he says that his theory is "so
thorough-going and consistent, that I cannot
now read other books on the subject without
indignation. ... One of the most important
axioms is, that as the quantity of any
commodity, for instance, plain food, which a
man has to consume, increases, so the utility
61
Journal & Proceedings of the Royal Society of New South Wales
Marks - WMam Stanley Jevons FPhSN 1856-59
or benefit derived from last portion used
decreases in degree.” (H, Jevons, 1886, p.
151). This is decreasing mar^al utility, as we
now know it.
Jevons wrote up his theory of value in a
paper for the British Association for the
Advancement of Science in 1862, which was
printed in 1866. There was little recognition
in Britain, given the sway of John Stuart Mills'
ideas. In his 1871 book he described his
theory in greater depth.
In establishing his utility theory, Jevons'
training in science and measurement was very
important. The quantities in the theory
(quantities, prices) could be exactly measured,
although its maximand, utility, was subjective.
“A unit of pleasure or of pain is difficult even
to conceive; but it is the amount of these
feelings which is continually prompting us to
buying and selling, borrowing and lending,
labouring and resting, producing and
consuming; and it is from the quantitative
effects of the feelings that we must estimate
their comparative amounts. We can no more
know or measure gravity in its own nature
than we can measure a feeling; but, just as we
measure gravity by its effects in the motion of
a pendulum, so we may estimate the equality
or inequality of feelings by the decisions of
tine human mind.” (fevons, 1871, p. 11),
For Jevons, value was directly a function
of utility. Perhaps I can illuminate how this
notion was new in 1871 by an anecdote from
1961. 1 had just turned 15, the age in Victoria
in those days when boys became men, at least
as far as the barber was concerned: we were
now, under the current price regulations,
charged men's prices for our haircuts. I
remember sitting in Mr. Merrknan's chair and
arguing with him that the prices should be
reversed: cutting men’s hair is easier than
cutting boys’ hair; men might begin to lose
their hair as they age; the conversation with
men is more interesting for the barber; and
men sit still without willing. So men’s
haircuts should cost less, young Robert
argued = in vain.
Turns out, although I didn’t know it, that
my argument was consistent with the classical
theory of value (from before 1860), which
states, roughly, tliat value, broadly speaking,
derives from the labour used to produce the
product, here the haircut: the cost to the
barber of a man’s hairciit is less than a boy’s,
so the price charged should be lower.^
I had overlooked a development dating
from Jevons’ writings of the 1860s, in which
he argued that the value (of the haircut)
depends on the utility the customer associates
with it. And men care more about their
appearance than do boys, and moreover are
able to pay more than boys can. So, on
Jevons’ theory of neoclassical value, men’s
haircuts should cost more than boys’ do, so
long as the barber is happy to cut at that
price.
From his theory of utility and value,
Jevons constructed a theory of exchange and
a theory of labour supply and capital.
Although he did not know it in 1862 or 1871,
his utility theory was not the first, which he
generously acknowledged in the preface to
the second edition of his book (in 18794^).
But many of his theoretical contributions,
developed independently, were ori^al,
unassailable, and of great usefulness.^
Jevons’ economic ideas continue to
resonate. Two of Ms applied studies received
much more attention at the time. His 1863
3 My argument also echoed the “labour theory of value,”
used in Karl Marx’s Capital (1867) in a misguided
attempt to explain relative prices. Marx has been
criticised for ignoring developments in economic theory,
such as those of Jevons’.
4 See http:/ / oIl.libert\Hnd.org/ titles /jevons-the-tlieoiv-
of-poHtical-economv
5 It must have been the Zeitgeist two other men, Carl
Menger (184(D 1921), an Austrian, and Leon Wakas
(1834D 1910), a Frenchman, published similar books in
the 1870s. These three are the pioneers of the
marginalist revolution of neo-classical microeconomics.
62
Journal & Proceedings of the Royal Society of New South Wales
Marks — William Stanley Jevons FPhSN 1856-59
pamphlet on the price of gold made a
remarkably accurate estimate of a 9 per cent
fall in the value of gold between 1848 and
1860, a time when many gold mines had
opened in California and Australia. In doing
so, he virtually invented the technique for
constructing price index numbers; John
Maynard Keynes, a first-rate mathematician,
stated that Jevons had “made as much
progress in this brief pamphlet as has been
made by aU succeeding authors put together.”
(Keynes, 1936, p. 525.)6
In his influential 1865 book. The Coal
Question^ Jevons was pessimistic about Great
Britain’s hiture as its coal resources — the
fuel of its industrial revolution — - became
economically exhausted. He argued that
increased “economy” of coal use, by which
he meant increased energy efficiency, would
not delay this date. He concluded:
It is wholly a confusion of ideas to
suppose that the economical use of fuel
is equivalent to a diminished
consumption. The very contrary is the
tmth.
That is, he was arguing that increased energy
efficiency “renders the employment of coal
more profitable, and thus the present demand
for coal is increased.”
This is the so-called Jevons paradox:
technological progress increases energy
efficiency (reducing the amount necessary for
any one use) but the rate of consumption of
that fuel might rise because of the lower
effective price and increased demand (the
rebound effect). Which of these effects
predominates continues to be a contentious
issue, and the debate has even escaped from
the pages of the learned journals. For
‘"Indeed, according to Harrod (1951, pp. 106), the 22-
year-old Keynes, on first reading Jevons’ work wrote, “I
am convinced that he was one of the minds of the
century.”
instance, in 2010, an article in the New Yorker
(Owen 2010) discussed the Jevons paradox,
and there was a debate in the pages of tlie
New York Times in 2012.^ Castles (2016)
overlooked this continuing influence of
Jevons on economic thinking, 150 years after
Coal was published.
In 1864 Jevons published a book based
on George Boole’s system of logic. In 1869
he built his “Logic Piano,” a device for
performing a function provided today by a
truth table. He had essentially mechanized
Boolean logic, a key aspect of contemporary
computing; he was the first. ^ (Barrett and
Connell, 2006). Later he became Professor of
Economics at University College, London. In
1872 he became only the second economist
to be elected as a Fellow of the Royal Society
of London. Flis son, Herbert Stanley Jevons,
published papers in this Journal in 1911 and
1912 on geology.
He was a true polymath. No other
Fellow of our Royal Society or its antecedents
(apart from Charles Darwin^) has had such an
impact on the intellectual life of the world.
Swimming in the English Channel on August
13, 1882, he drowned, aged 46. We should
honour his memory.
Acknowledgements
I’d like to acknowledge assistance from Geoff
Harcourt, Eve Wynhausen, Chris Adam, and
Raja Junankar.
References
Anon., 2012, The siren song of energy efficiency.
The New York Times, March 19.
Indeed, there are over 2,200 articles/books in Google
Scholar that include the phrase “Jevons Paradox.”
^ The original Jevons’ Logic Machine is held at the
Museum for the History of Science, Oxford.
^ Kelly (2009) includes a copy of a letter firom Darwin of
Oct. 28, 1879, accepting his honorary membership of
the Royal Society.
63
JouRN.\L & Proceedings of the Royal Society of New South Wales
Marks - William Stanley Jevons FPhSN 1856-59
Barrett, Lindsay, and Connell, Matthew, 2005,
Jevons and the Logic Tiano,’ The Tutherford
Journal 1, No. 1, December
http:/ /v^vi^.rudierfordi ournal.org/ artideOlOlOS
.hanl
Black, R. D. CoUison (ed.), 1973, Tapers and
Correspondence of William Stanley Jevons, Volume IJ
Correspondence, 1850-1862, London: Macmillan,
for the Royal Economic Society.
Bourke, Iris, 1955, Australia’s first pictoriakst,
Australian Photo Review, pp. 6— 23, Jan.
Castles, Ian, 2016, The curious economist: William
Stanley Jevons in Sydnty, Journal and Proceedings of
the Royal Society ofN.S.W., this issue.
Harrod, Roy, The life of John Maynard Kynes.
London: Macmillan. 1951.
Jevons, Harriet A, (ed.), 1886, letters and journal of
W. Stanley Jevons, London: Macmillan.
https:/ / arcliive.org/ stream /letters] ournalofOOie
voiala#page/ n7 / mode/ 2up
Jevons, William Stanley, 1859, The climatology of
Australasia, nine chapters m Waugh's Australian
Almanac 1859. Sydney: James William Waugh.
Jevons, William Stanley, 1863, A serious fall in the
value of gold ascertained, and its social effects set forth,
London: Edward Stanford.
https:/ / arcliive.org/ stteam/ aseriousfaUinvOOiev
ogoog
Jevons, William Stanley, 1862, Notice of a general
mathematical theory of political economy,
presented to the British Association for the
Advance of Science, Cambridge, October.
Jevons, William Stanley, 1866, Brief account of
a general mathematical theory of political
economy, Journal of the Royal Statistical Society, 29:
282-87, June.
http:/ /la.utexas.edu/ users /hcleaver/ 368/ 368iev
onsmathecotable.pdf
Jevons, William Stanley, 1866, The coal question: An
inquiry concerning the progress of the national and the
probable exhaustion of our coal-mines. (2nd. ed.)
London: Macmillan.
http:/ / w^wvceconlib.org/librar)AYPDBooks/Je
vons / ivnCQCover.html
Jevons, William Stanley, 1871, The theory of political
economy, first edition, London: Macmillan.
Kelly, Jak, 2009, Editorial: A well known member.
Journal and Proceedings of the Royal Society ofM.S.W.
142(1-2): 1-3.
Keynes, John Maynard, 1936, William Stanley
Jevons, 1835—1882, a centenary allocution on his
life and work as economist and statistician,
Journal of the Rpyal Statistical Society, 99: 516- 555.
La Nau2e, John A., 1953, The concept of Jevons’
utility theory. Economica, 20: 356—8, Nov. Marx,
Karl, 1867, Capital: critique of political economy,
Vol 1 (first English edition, 1886).
Owen, David, 2010, The efficiency dilemma. The
New Yorker, Dec. 20 and 27.
http:/ / www.newt^orker.com/ maga2ine/201 0/ 1
2/20/ the-efficiency-dilemma
Vaughn, Kelly, 2012, Jevons paradox: the debate
that just won’t die, RMl Outlet, March 20.
http://blog.mii.org/blog Tevons Paradox
64
Journal & Proceedings of the Royal Society of New South Wales, vol. 1 49, parts 1 & 2, 20 1 6,
pp. 65-82. ISSN 0033-9173/16/010065-18
The evolutionary history of flowering plants
Charles S.P. Fosteri
1 School of Life and Environmental Sciences, University of Sydney, New South Wales 2006, Australia
This paper was an RSNSW Scholarship Winner in 2015
Email: charles.foster@sydney.edu.au
Abstract
In terms of species richness and important ecological roles, there are fev/ biological groups
that rival the success of flotvering plants (Angiospermae). Angiosperm evolution has long
been a topic of interest, with many attempts to clarify their phylogenetic relationships and
timescale of evolution. However, despite this attention there remain many unsolved questions
surrounding how and when flowers first appeared, and much of the angiosperm diversity
remains to be quantified. Here, I review the evolutionary history of angiosperms, and how
our understanding of this has changed over time. I begin by summarising the incredible
morphological and genetic diversity of flowering plants, and the ways in which this can be
studied using phylogenetic inference. I continue by discussing both the relationships between
angiosperms and the other major lineages of seed plants, and the relationships between the
main groups within angiosperms. In both cases, I outline how our knowledge has changed
over time based on factors such as the different conclusions drawn from morphological and
genetic data. I then discuss attempts to estimate the timescale of angiosperm evolution and
the difficulties of doing so, including the apparent conflict between ages derived from fossil
and molecular evidence. Finally, I propose future directions for angiosperm research to help
clarify the evolutionary history of one of the most important groups of organisms on the
planet.
Introduction
he diversity and interactions of life on
Earth have long been of scientific inter¬
est. Quantifying biodiversity and the times¬
cale over which it arose allows inferences
about the biological history of the planet
to be made, and can provide insight into
how ecosystems might change in response
to events such as cHmate change (ThuiUer
et al. 2011; Bellard et al. 2012). Flowering
plants (angiosperms) have been of particular
focus because of their important economic
and cultural roles within society, as well as
their ubiquity and importance within natu¬
ral ecosystems. Specifically, angiosperms
sequester large amounts of carbon from
the atmosphere, and act as primary produc¬
ers of food for many animal groups, with
their spread and appearance shaping habitat
strucmre globally (Brodribb and Feild 2010;
MagaUon 2014). In addition, angiosperms
have developed important mutualistic rela¬
tionships with many groups of organisms,
such as pollination interactions with insects,
birds, and small mammals (van der Niet and
Johnson 2012; Rosas-Guerrero et al. 2014).
However, to properly quantify the extent
and impact of groups such as angiosperms,
biological entities must first be recog¬
nised and described into distinct groups
such as species, and, ideally, placed into
higher-order classifications. The goal is to
65
JouRN.\L & Proceedings of the Royal Society of New South Wales
Foster — Piistory of Flowering Plants
recognise groups that contain only the
descendants of a common evolutionary
ancestor (monophyletic groups), which
represent natural evolutionary groups.
For most of history, biological groups
and the relationships between them have
been recognised through observations of the
form and stmcture of organisms. When
these data are shared between two or more
taxa after being inherited from their most
recent common ancestor, they are known as
synapomorphies. In addition to aiding the
classification of extant taxa, these
morphological data are also able to link
extant and extinct diversity through
comparison with the fossil record, which can
suggest a timescale of evolution. However,
morphological data often cannot reliably
distinguish between competing taxonomic
h}podieses because of a lack of informative
characters, or can be misled by the
independent evolution of similar traits in
organisms that are not closely related
(convergent evolution). Morphological data
have been supplemented by molecular data
since the inception of molecular
phylogenetics in the mid-20^^ Century.
Molecular data typically comprise
sequences of the nucleotides of DNA, or the
amino acids that they encode. Each
nucleotide or amino acid within a sequence
represents a character that can be used for
phylogenetic analysis. Therefore, molecular
data sets can contain millions of characters
for phylogenetic reconstruction, which
makes such data sets especially usefial for
evaluating the taxonomic hypotheses that
have been suggested by morphology.
Analysis of molecular data is also useful for
estimating the evolutionary timescale of
organisms using molecular clocks (Lee and
Ho 2016), especially for groups with poor
fossil records.
Both morphological and molecular data
have been used extensively to evaluate the
diversity of angiosperms. Angiosperms are
among the most species-rich groups of
organisms on the planet, and are by far the
largest group of plants. The exact number of
species is difficult to determine because of
high amounts of taxonomic synonymy, and
the fact that many species potentially remain
to be discovered (Bebber et al. 2010; Pimm
and Joppa 2015). Despite this, we can be
fairly certain that there are at least 350,000
species of angiosperms, and probably c.
400,000 in total (Pimm and Joppa 2015). As
expected in a group of this sEe, there is
extreme variation in morphology, life history
characteristics, and growth form.
Angiosperms variously exist as herbaceous
annuals, vines, lianas, shrubs or trees, and
can be found growing in aquatic or terrestrial
environments, or even growing on and/or
parasitising other plants.
Similarly, there is large variation in
genome size and content withiu
angiosperms. For example, it is estimated
that throughout their evolutionary history
over 70% of angiosperms have had an
increase in the number of copies of
chromosomes contained within each cell
(ploidy level) from the typical diploid state
(Levin 2002), Most of the fiinctions essential
for growth and development are controlled
by genes located within the cell nucleus,
which are collectively known as the nuclear
genome. Vans japonica Franch., a small
herbaceous plant native to Japan, has the
largest accurately measured genome known
to science (Pellicer et al. 2010). At nearly 150
billion nucleotides, its octoploid genome is
more than 50 times larger than the human
genome, and nearly 2500 times larger than
the smallest known plant nuclear genome of
Genlisea tuherosa Rivadavia, Gonella &
A.Fleischm., a carnivorous angiosperm from
Brazil (Fleischmann et al. 2014).
Plant cells also contain specialised
organelles known as chloroplasts and
66
Journal & Proceedings of the Royal Society of New South Wales
Foster - History of Flowering Plants
mitochondria, which are responsible for the
essential processes of photosynthesis and
cellular respiration, respectively. Both of
these organelles are predominantly
uniparentaUy inherited and contain thek own
independent genomes, which is thought to
be because of thek origins as free-living
organisms that were engulfed by early
eukaryotic cells in separate endosymbiotic
events (Sagan 1967; Schwartz and Dayhoff
1978). The chloroplast genome varies
substantially among angiosperms, with the
order of genes differing between groups, and
with some genes being lost completely. For
example, the chloroplast genome is
drastically reduced in many parasitic plants,
with many genes important for
photosynthesis having been lost (Bungard
2004).
The mitochondrial genome of plants is
more enigmatic, and is disproportionally less
studied than the nuclear and chloroplast
genomes. Plant mitochondrial genomes are
large compared with animal mitochondrial
genomes, and thek content is highly
dynamic, with many gene gains, losses,
transfers, duplications and rearrangements,
as weU as a large proportion of repeated
elements and introns (Kitazaki and Kubo
2010; Galtier 2011). Of dkect importance
for reconstmcting the evolutionary history of
plants is that the three genomes evolve at
very different rates. The nuclear genome
evolves at the highest rate, the chloroplast
genome evolves at an intermediate rate, and,
in contrast to its dynamic nature, the
mitochondrial genome has by far the lowest
evolutionary rate (Wolfe et al. 1987).
The global dominance of angiosperms
indicates that they are ideally adapted to exist
within many different habitats, and thek
great morphological and genomic variation
suggests a history of varied selective
pressures. This has long challenged those
who have sought to quantify how such a
diverse group arose over a supposedly short
period of time. Indeed, the traditional view
is that angiosperms originated in the early
Cretaceous. The subsequent appearance of
fossils with highly diverse morphologies,
over what was apparently an extremely rapid
timescale, was famously described by Darwin
as an “abominable mystery” in a letter to
Joseph Hooker in 1879 (first published in
Darwin and Seward 1903).
To understand fully the evolutionary
history of angiosperms, thek diversity needs
to be characterised in a phylogenetic context.
This approach indicates whether key traits
for success are clade-specific, or have
evolved multiple times in parallel.
Additionally, incorporating temporal
information into these analyses can allow
inferences to be made about the
envkonmental conditions that might have
driven angio sperm diversification.
In this review, I begin by discussing our
understanding of the relationships among
the major seed plant Kneages, and the
importance of this for reconstructing the
origin of flowers. I then discuss the
relationships of the major lineages within
Angiospermae, and examine estimates of the
evolutionary timescale of angiosperms. I
propose a number of the future dkections
that are likely to improve our understanding
of the evolutionary history of angiosperms.
Higher relationships of angiosperms
and the origin of flowers
Angiosperms are recognised as members of
the superdivision Spermatophyta along with
cycads, conifers, gnetophytes, and Ginkgo.
The last four extant cone-bearing lineages are
known as acrogymnosperms, whereas extant
and extinct cone-bearing lineages combined
are known as gymnosperms (Cantino et al.
2007). The five extant spermatophyte
lineages are linked by the production of
67
Journal & Proceedings of the Royal Society of New South Wales
Foster — History of Flowering Plants
seeds. Estimates of the number of seed
plant species vary, but are consistently in the
region of many hundred thousand species
(Govaerts 2001; Scotland and Wortley 2003).
Among other potential factors, the success
of these lineages is perhaps due to the
diversification of regulatory genes important
for seed and floral development following
ancient whole-genome duplication events
along the lineages leading to seed plants and
angiosperms (Jiao et al. 2011).
Angiosperms can be readily
distinguished from gymnosperms through a
suite of synapomorphies. These include the
presence of flowers with at least one carpel,
which develop into fruit (c£ the “naked”
seeds of gymnosperms); stamens with two
pairs of poUen sacs (cf. the larger, heavier
corresponding organs of gymnosperms); a
range of features of gametophyte structure
and development, including drastically
reduced male and female gametophytes
compared with gymnosperms; and phloem
tissue with sieve tubes and companion cells
(cf. sieve cells without companion cells in
gymnosperms) (Doyle and Donoghue 1986;
Soltis and Soltis 2004). The production of
endosperm through double fertilisation was
previously considered to be a further
synapomorphy of angiosperms, but this
phenomenon has also been observed in
some gnetophyte lineages (Friedman 1992;
Carmichael and Friedman 1996).
Collectively, the synapomorphies of
angiosperms are thought to be responsible
for providing the evolutionary advantages
that led to theit global dominance, which
coincided with a decline in gymnosperm
diversity (Bond 1989). However, to
reconstruct the evolution of these characters
and evaluate their importance for
angiosperm evolution, it is necessary to
determine which lineage of seed plants is
most closely related to angiosperms. The
majority of earlier studies focused on
evaluating the seed plant phylogeny,
including determining the sister lineage to
angiosperms, using comparative morphology
to assess homology of the reproductive and
vegetative structures of the seed plant
lineages (e.g., Doyle and Donoghue 1986).
One major hope was that determining
the sister lineage to angiosperms might prove
especially useful for inferring the origin and
structure of the first flowers. Throughout
the 20* cenmry, the two main hypotheses
for the origin of flowers were that they
evolved from branched, unisexual
reproductive structures found in most
gymnosperms ("pseudanthial" theory,
Wettstein 1907), or that flowers evolved
from bisexual, flower-like structures, such as
in the extinct group Bennettitales
(’'euanthial” theory, Arber and Parkin 1907).
The inferred homology of morphological
structures consistently su^ested that
gnetophytes were the extant sister lineage to
angiosperms, with several potential close
(non-angiosperm) fossil relatives.
Specifically, various features of wood
anatomy and flower-Hke structures seemed
to suggest a close relationship between
angiosperms, gnetophytes, and the extinct
order Bennettitales, with this group being the
sister lineage to the rest of the gymnosperms
(Crane 1985; Doyle and Donoghue 1986).
Therefore, based on the strength of
morphological evidence, the euanthial theory
was the most popular view in the 20*
Century.
The acceptance of the euanthial theory,
coupled with the predominance of
Cretaceous Magnolia-\^^ fossils at the time,
led to suggestions that the ancestral flowers
were similar to present-day magnolias. This
implies that magnolias and their close
relatives were some of the earliest-diverging
angiosperm lineages (Endress 1987).
However, most molecular phylogenetic
studies from the 1990s onwards have
68
Journal & Proceedings of the Royal Society of New South Wales
Foster — Plistory of Flowering Plants
recovered different relationships between the
extant seed plant lineages. The dominant
theme in these modern studies is that all
extant gymnosperm lineages form a
monophyletic sister group to angiosperms
(Chaw et al. 1997; Bowe et al. 2000; Chaw et
al. 2000; Ruhfel et al. 2014; Wickett et al.
2014) (Figure 1). Particularly strong evidence
has emerged for a close relationship between
gnetophytes and conifers (Qiu et al. 1999;
Winter et al. 1999). Indeed, the evidence
seems to surest that gnetophytes might
even be nested within conifers and the sister
group to Pinaceae (Bowe et al. 2000; Chaw et
al 2000; Zhong et al 2010).
Overall, because none of the extant
gymnosperm lineages is more closely related
to angiosperms than to other gymnosperms,
they cannot directly inform hypotheses on
the homologies of angiosperm characters, or
on the sequence of development of these
characters (Doyle 2012). Therefore, while
the relationships among the major seed plant
lineages have been largely resolved, the
structural origin of flowers, and the affinity
of the earliest flowers to modern species,
remains controversial. Progress in this area
is likely to be achieved through improved
understanding of the relationships among
the major angiosperm groups.
Major relationships within
Angiospermae
The major relationships within angiosperms
have historically proved difficult to
determine, and have long been in a state of
flux. This has largely been due to differing
ideas of the characters, initially
morphological but later molecular, needed to
reconstmct the angiosperm phylogeny. An
early discovery was that flowering plants
have either one or two embryonic leaves
(Ray 1686""1704). While John Ray was the
first to observe this dichotomy, he later
followed Marcello Malpighi in referring to
these leaves as ‘cotyledons’. Accordingly,
flowering plants with one cotyledon have
subsequently been referred to as
monocotyledons or ‘monocots’, and those
with two cotyledons have been called
dicotyledons or ‘dicots’.
Although the most widely known early
classification scheme by Linnaeus was based
solely on floral reproductive characters, the
division into monocots and dicots has since
been recognised as an important diagnostic
feature to inform classification, with varying
implications for the angiosperm phylogeny.
A minority of early authors argued that some
key morphological differences between
monocots and dicots, such as vascular
bundle anatomy, were irreconcilable with a
monophyletic origin of angiosperms.
Instead, these authors argued that
angiosperms should be recognised as a
polyphyletic group (= derived from more
than one common evolutionary ancestor)
(e.g, Meeuse 1972; Krassilov 1977).
However, the predominant view was that
angiosperms are monophyletic, and the
division into monocots and dicots
constitutes a natural split within flowering
plants. This was echoed in many angiosperm
classification systems developed in the 20^^
century, including the highly influential
Takhtajan (1980) and Cronquist (1981)
systems.
To infer the evolutionary relationships
within monocots and dicots, many cladistic
analyses were undertaken in the latter half of
the 20* century using pollen, floral, and
vegetative characters. This approach led to
many informal subgroups being proposed.
For example, Donoghue and Doyle (1989b)
recognised five major groups of
angiosperms, corresponding to MagnoHales,
Laurales, Winteraceae-like plants,
‘paleoherbs’ (‘primitive’ herbaceous lineages
including water lilies and A.mhorelld)^ and
69
JouRN.\L & Proceedings of the Royal Society of New South Wales
Foster ~ History of Flowering Plants
Figure 1: The relationships among seed plant lineages, scaled to geological time based on fossil
ages. Numbers in green circles refer to the following: (1) oldest Ginkgo fossil (Yang et al 2008);
(2) oldest cycad fossil (Gao and Thomas 1989); (3) oldest gnetophyte fossil (Rydin et al. 2006);
(4) oldest conifer fossils (Wieland 1935); (5) oldest angiosperm fossils (discussed in Doyle
2012); (6) oldest acrogymnosperm fossil ; (7) an estimated maximum age for crown-group seed
plants (discussed in MagaUon and CastiUo 2009; Foster et al. 2016).
plants with tricolpate pollen. Although the
constituent members of the subgroups
varied across studies, the recognition of
tricolpates as a monophyletic group was a
consistent finding (e.g., Donoghue and
Doyle 1989b; Donoghue and Doyle 1989a),
leading to suggestions that dicots had
multiple evolutionary origins (Endress et al.
2000; Endress 2002). Indeed, stratigraphical
studies in which triaperturate pollen
(tricolpate) fossils were consistently found to
originate in younger sediments than both
monocots and non-tricolpate dicots had
already hinted that dicots did not form a
monophyletic group (Doyle 1969).
Consequently, Doyle and Hotton (1991)
chose to recognise tricolpates as distinct
from the rest of the dicots, coining the term
‘eudicots’ for this group.
Taxonomic concepts for the major
angiosperm groups have changed over time,
which makes it difficult to chronicle
concisely the changing opinions about the
earliest-diverging angiosperms. For example,
the group Magnoliidae now has a very
different circumscription compared with the
past, so statements in earlier studies
regarding the relationships between
70
Journal & Proceedings of the Royal Society of New South Wales
Foster — IListory of Flowering Plants
magnoliids and other groups might no
longer be applicable. Nevertheless, it is clear
that the most common view historically was
that Magnolia-like flowers probably occupied
a position at or near the root of the
angiosperm phylogeny. However, there
were other suggestions for the earliest-
diverging angiosperm lineages, including
Piperales+Chloranthales, several of the
lineages in the formerly recognised paleoherb
group, or even monocots (Burger 1977,
1981).
Attempts to clarify the relationships
within the angiosperm phylogeny have since
been gready strengthened by the inclusion of
molecular data. Some aspects of early
classification schemes based on morphology
have been strongly supported by molecular
data (reviewed by Endress et al. 2000;
Endress 2002). For example, the key
concepts of the monophyly of angiosperms,
monocots and eudicots, the polyphyly of
dicots, and the position of magnoHids as an
early diverging angiosperm lineage, were all
further supported by molecular data
(Endress et al. 2000). However, many
molecular estimates of angiosperm
evolutionary relationships have contradicted
estimates based on morphological data. For
example, molecular data have firmly resolved
the family Hydatellaceae within
Nymphaeales, rather than within Poales as
former morphology-based studies had
concluded (Saarela et al. 2007). Molecular
data have also helped to clarify the extent of
convergent evolution within angiosperms,
such as C4 photosynthesis evolving
independently at least 60 times (Sage et al.
2011).
Arguably the most important finding
from analyses of molecular data has been the
rooting of the angiosperm phylogeny.
Success was not immediate, with
disagreements being found among the results
of molecular analyses, depending on the
choice of molecular markers. An influential
early attempt with molecular data to resolve
the seed plant phylogeny and, necessarily, to
determine the earHest-diverging angiosperm
Hneage, analysed sequences for the
chloroplast rkL gene from nearly 500 seed
plant taxa using maximum parsimony (Chase
et al. 1993). In this case, the widespread
aquatic genus Ceratophjllum was found to be
the sister lineage to all other flowering plants.
However, this has subsequently been found
to be an anomalous result seemingly unique
to single-gene parsimony analyses of rAL. A
series of studies in 1999 found that the
monotypic genus Mmhorella is strongly
supported as being the sister lineage to all
other flowering plants (Mathews and
Donoghue 1999; Parkinson et al. 1999; Qiu
et al. 1999; Soltis et al. 1999), and this finding
has subsequently been supported by nearly
aU large multigene analyses (Moore et al.
2007; Soltis et al. 2011; but see Goremykin et
al. 2013; Ruhfel et al. 2014; Wickett et al.
2014; Xi et al. 2014; Goremykin et al. 2015).
These studies have also revealed that the
base of the angiosperm phylogeny
constitutes a grade of several successive
lineages, originally referred to as the ANITA
(M/;^^or^/Z:?/Nymphaeale s / lUiciaceae-
"kv^mem'Aeem-Austrohaikjd) grade, but now
known as the ANA
(M/^/?^?r(?/Z:7/Nymphaeales/Austobaileyales)
grade.
The remaining -99.95% of angiosperms
are collectively referred to as
Mesangiospermae. Within this group, five
major Hneages are recognised: Chloranthales,
MagnoHidae, CeratophyUales, monocots, and
eudicots (clade names here are standardised
to Cantino et al. 2007). Unfortunately,
despite large increases in the amount of
available genetic data and improved analytical
techniques, the relationships among these
mesangiosperm groups have remained
uncertain (Figure 2). When analysing
71
Journal & Proceedings of the Royal Society of New South Wales
Foster = History of Flowering Plants
chloroplast genome sequences, the most
common finding is that eudicots
+Ceratophjlkm form the sister group to
monocots, with these three lineages being
the sister group to magnoMds
TChloranthales. Large nuclear DNA data
sets, which have only become available in
recent years, tend to resolve different
relationships. For example, they have
supported a sister relationship between
eudicots and magnoliids+Chloranthales, with
monocots being the sister group to these
three lineages (Wickett et al. 2014).
However, the number and choice of nuclear
DNA markers can affect inferred
relationships within Mesangiospermae. For
example, analysis of a selection of 59 low=
copy nuclear genes inferred a grouping of
Ceratophjlkm+ChlomnAi'Ae.s and eudicots,
with successive sister relationships to
magnoliids and monocots (Zeng et al. 2014),
Additionally, the choice of phylogeny
reconstruction method can lead to the
estimation of different topologies (Xi et al.
2014).
Nevertheless, despite conflicting
topologies sometimes being inferred, we
currently have an understanding of tine
angiosperm phylogeny that is greater than at
any other time in history. The power of
molecular data to resolve the historically
challenging relationships among flowering
plants is now well established. In response
to the rapid advances in the field, a
cosmopolitan consortium of researchers
regularly collaborate to release timely
summaries of the state of knowledge of the
angiosperm phylogeny (see Angiosperm
Phylogeny Group 1998, 2003, 2009, 2016).
We now have a viable framework to allow
fields related to phylogenetics to flourish and
provide a greater understanding of the
important evolutionary steps that have
contributed to the overwhelming success of
angiosperms, such as through evolutionary
developmental biology (evo^devo) studies
(Preston and Hileman 2009). However, to
gain a fuller understanding of the
evolutionary history of angiosperms, it is
necessary to know more than just the
relationships among the major flowering
plant groups; a reliable estimate of the
angiosperm evolutionary timescale is also
needed.
Evolutionary timescale of
angiosperms
To understand how an^osperms came to
dominance, including how the crucial
morpholo^cal traits that led to their success
first evolved, it is necessary to have some
idea of the timescale of angiosperm
evolution. Traditionally, the evolutionary
timescale of organisms has been elucidated
through study of the fossil record. In this
approach, the first appearance of each taxon
in the fossil record, as determined by
morphology, provides an indication of when
it first evolved. When considering the fossil
record, it is important to distinguish between
“crown” and “stem” groups. A crown group
is the least indusive monophyletic group that
contains aU extant members of a clade, as
well as any extinct lineages that diverged after
the most recent common ancestor of the
dade (MagaMon and Sanderson, 2001). In
contrast, a stem group is the most indusive
monophyletic group that contains aU extant
members of a dade, as well as any extinct
lineages that diverged from the lineage
leading to the crown group (Magallon and
Sanderson, 2001),
The fossil record of seed plants is
ancient, with the oldest fossils of
progymnosperms occurring in sediments
from the Late Devonian, ^365 mUHon years
ago (Ma) (Fairon-Demaret and Schedder
1987; Rothwell et al, 1989; Fairon-Demaret
1996).The fossil record of gymnosperms
72
Journal & Proceedings of the Royal Society of New South Wales
Foster — History of Flowering Plants
Eudicots
Ceratophyllales
Monocots
Magnoliids
Chloranthales
ANA
(d)
Eudicots
Monocots
Magnoliids
Ceratophyllales
Chloranthales
ANA
cpDNA
Morphology
+ cpDNA
Monocots
Magnoliids
Eudicots
Ceratophyllales
Chloranthales
ANA
Figure 2: A comparison of several different estimates of the relationships among eudicots,
magnoliids, monocots, Ceratophyllum, Chloranthales, and ANA-grade angiosperms, based
on the comparison presented in Zeng et al. (2014), The different topologies represent
findings from studies using nuclear DNA (nrDNA), chloroplast DNA (cpDNA),
mitochondrial DNA (mtDNA), and a combination of morphological and molecular data.
A sample of suitable references for the topologies are as follows: (a) Zhang et al. (2012); (b)
Moore et al. (2011); Zeng et al. (2014); (c) Moore et al. (2007); Moore et al. (2010); Foster et
al. (2016); (d) Qiu et al. (2010); (e) Endress and Doyle (2009).
is rich, with fossils becoming common from
the Late Carboniferous to Early Triassic
(MagaUon 2014), and reveaHng an extinct
diversity far greater than the extant diversity.
The oldest known fossil that can be
confidently assigned to the stem group of
angiosperms has suggested that angiosperms
arose as early as 247.2-242.0 Ma (million
years ago) (HochuH and Feist-Burkhardt
2013). Accepted poUen fossils (microfossils)
suggest that crown-group angiosperms first
Unfortunately,, the fossil record of
angiosperms is not as extensive or
informative.
appeared in the Valanginian to early
Hauterivian (early Cretaceous, -139.8-129.4
Ma), albeit ia sparse amounts, with vast
amounts of angiospermous microfossils
occurring by the Barremian (—129.4-125
Ma) (Doyle 2012). There is a noticeable
disparity in the number and presence of
73
Journal & Proceedings of the Royal Society of New South Wales
Foster — History of Flowering Plants
fossils between lineages, particularly at the
family level and below, with many excellent
fossils being present for some groups but
none for others (Magalon 2014).
While fossil data have traditionally
provided the only source of information
about the evolutionary timescale of major
groups, molecular dating techniques provide
a compelling alternative, especially for groups
that lack fossils. In these approaches,
evolutionary timescales can be estimated
using phylogenetic methods based on
molecular clocks. When the concept of the
molecular clock was &st proposed,
evolutionary change was assumed to
correlate linearly with time and to remain
constant across lineages (‘'strict” molecular
clock) (Zuckerkandl and Pauling 1962).
However, it has since become dear that
strictly clocklike evolution is the exception,
rather than the rule (Welch and Bromham
2005).
Rates of molecular evolution vary
substantially across vascular plant lineages
(Soltis et al. 2002), and are often strongly
correlated with Mfe history strategies. For
example, substitution rates in herbaceous
annual lineages of angiosperms are known to
be substantially higher than in woody
perennial plants (Smith and Donoghue 2008;
Lanfear et al. 2013). Consequently, a variety
of molecular clock models have been
developed to account for evolutionary rate
variation among lineages (Ho and Duchene
2014). Fossil data are still intricately linked
with these methods, because fossils are used
to provide temporal information to calibrate
the molecular dock, thereby providing
absolute rather than relative ages of nodes.
For example, in Bayesian analyses, temporal
information is incorporated through
calibrations priors, which can take the form
of a variety of probability distributions (Ho
and Phillips 2009). In the absence of fossils
for a particular group being studied,
biogeographic events and rate estimates from
other groups can be used as calibrations, but
these are subject to a wide range of errors
(Ho et al. 2015).
Collectively, molecular dating studies
have yielded remarkably disparate estimates
for the age of crown-group angiosperms
(summarised in Bell et al 2010; Magallon
2014; Foster et al. 2016). Inferred ages have
ranged from the extreme values of 86 Ma
(when considering only the 3rd codon
positions of r^rL; Sanderson and Doyle
2001) to 332.6 Ma (Soltis et al. 2002). Most
age estimates fall between 140 and 240 Ma,
but this still represents a substantial amount
of variation. Additionally, the earliest
analyses found that crown-group
angiosperms were considerably older than
impKed by the fossil record, in some cases by
more than 100 million years (e.g. Martin et al.
1989). Smaller disparities between molecular
and fossil estimates were obtained in later
studies (e.g. Sanderson and Doyle 2001).
However, some more recent estimates have
tended to support a more protracted
timescale for angiosperm evolution (e.g.
Smith et al. 2010), echoing the results of the
earliest molecular studies.
Progress in molecular dating can be
characterised in terms of increasing
methodological complexity and improving
sampling of taxa and genes (Ho 2014). A
persistent problem, however, has been the
need for a trade-off between taxon sampling
and gene sampling. Low gene sampling has
been typical of studies of an^osperm
evolution, albeit with some other exceptions,
including the 12 mitochondrial genes
analysed by Laroche et al. (1995), 58
chloroplast genes analysed by Goremykin et
al. (1997), 61 chloroplast genes analysed by
Moore et al. (2007), and the 83 chloroplast
genes analysed by Moore et al. (2010).
However, most of these studies had sparse
angiosperm taxon sampling. Among the few
74
Journal & Proceedings of the Royal Society of New South Wales
Foster -- History of Flowering Plants
other studies that have included more than
50 taxa, the largest number of genes sampled
was five. The largest taxon samples have
been those of Zanne et al. (2014), which
used a sta^ering 32,223 species, and
MagaUon et al. (2015), which included 792
angiosperm taxa and one of the largest
samples of fossil calibration points ever used.
An exception to the above trade-off between
taxon and gene sampling is the study by
Foster et al. (2016), which analysed 76
chloroplast genes from 193 angiosperm taxa.
The most controversial aspect of
angiosperm molecular dating studies has
been an apparent incongruence between
molecular estimates and those extrapolated
purely from fossil occurrence data. Many
modern molecular dating estimates without
strongly informative temporal calibrations
tend to surest that angiosperms arose in the
early to mid-Triassic (Figure 3) (Foster et al.
2016), which impHes a considerable gap in
the fossil record (Doyle 2012). This
contradicts the claim that the evolutionary
history of crown-group angiosperms is well
represented in the fossil record (MagaUon
2014), despite several lines of evidence
supporting this suggestion: the gradual
increase in abundance, diversity, and
distribution of fossU angiosperms; the
ordered progression of both morphological
and functional diversification; and the
agreement between the stratigraphic record
and molecular data in the sequential
appearance of angiosperm lineages.
If the fault Hes instead with the molecular
estimates, then it has been su^ested that the
substantial disparity between molecular and
fossU-based estimates of the age of crown
angiosperms might be a result of the choices
of molecular markers, taxa, caUbrations, or
models of rate variation (MagaUon 2014).
Particular blame has been placed on the
inabUity of molecular dating methods to
account properly for non-representative
sampling of angiosperms and Ufe history-
associated rate heterogeneity (BeauHeu et al.
2015) . However, comprehensive
investigations of the impact of models,
priors, and gene sampling on Bayesian
estimates of the angiosperm evolutionary
timescale, using a genome-scale data set and
numerous, widely distributed fossU
caUbrations, have stiU yielded remarkably
robust estimates of a Triassic origin of
angiosperms (Foster et al. 2016). This
impUes a long period of no angiosperm
fossUisation, or that fossUs of this age simply
remain to be discovered (but see Wang et
al2007; Gang et al. 2016).
Despite the disparate estimates for
the origin of crown-group angiosperms, the
timescale of evolution within this group is
beginning to be understood with increased
precision. Of particular note is that estimates
for the origin of most modern angiosperm
orders seem to be consistent regardless of
tlie age inferred for the angiosperm crown
group (MagaUon et al. 2015; Foster et al.
2016) . Ordinal diversification is most
commonly estimated to have begun in the
early Cretaceous, and is concentrated
predominantly from this time through to the
mid-Cretaceous (MagaUon et al. 2015; Foster
et al. 2016). Modern angiosperm famiUes are
estimated to have originated steadUy from
the early Cretaceous, with the peak of famUy
genesis occurring from the late Cretaceous to
the early Paleogene (MagaUon et al. 2015)
During this time, the supercontinent Pangaea
largely completed its breakup into the
continents of the present day. Concurrently,
there were dramatic shifts in cHmate, with
global temperatures and CO2 levels far
higher than in the present day (Hay and
Floegel 2012), These changes, particularly in
temperature, would have had significant
impacts on the levels and efficiency of.
75
Journal & Proceedings of the Royal Society of New South Wales
Foster — History of Flowering Plants
Figure 3: A recent estimate of the angiosperm evolutionary timescale^ modified from Foster et
al. (2016). Numbers in parentheses after taxon names refer to the number of taxa included
from those groups in the study. Green circles indicate estimates of the crown age for lineages
when more than one taxon has been included, and the blue star indicates the inferred age for
the origin of crown-group an^osperms. The dashed line indicates the time by which aU
modern orders were inferred to have arisen.
photosynthesis (ElHs 2010; Hay and Floegel
2012). Selective pressures would have been
high, ultimately influencing the evolution of
an^osperms and, presumably, other taxa that
interacted with them.
Concluding remarks and future
directions
The substantial diversity and global
dominance of flowering plants have puzzled
and intrigued many researchers throughout
history. The classification of angiosperms
has long proved difficult because of the
monumental size and such varied
morphologies within this group.
Subsequently, the key evolutionary
innovations that first occurred to produce
flowers, as well as the reasons for the
overwhelming success of angiosperms, have
historically been obscured. Therefore, it is
reasonable to surmise that for most of
histojy, the relationship of angiosperms to
other seed plants, the relationships within
an^osperms, the timescale of angiosperm
evolution, and the reasons for the relative
success of an^osperms compared to
gymnosperms were all largely unknown or
not understood.
Thankfully, we have now made great
progress in the quest to answer these
questions. Work remains to identify
potential stem-group relatives of seed plants,
but we now have reliable estimates of the
phylogeny of extant seed plants. However,
the most widely accepted seed plant
phylogeny su^ests that no extant
gymnosperm lineage preserves the
evolutionary steps that led to the origin of
the first flowers. Therefore, in some respects
the resolution of the seed plant phylogeny
has been somewhat of a disappointment for
those wanting to reconstruct the
76
Journal & Proceedings of the Royal Society of New South Wales
Foster — Fiistory of Flowering Plants
development of the flower (Doyle 2012).
While this might be considered a setback,
our gready improved knowledge of the
angiosperm phylogeny, including a strongly
supported position for the root, allows
increasingly sophisticated questions to be
asked about angiosperm macroevolution
(e.g, Turcotte et al. 2014; Zanne et al. 2014).
Similarly, our modern estimates for the
timescale of angiosperm evolution allow us
to explore hirther the selective pressures that
might have shaped the present-day
distribution and diversity of flowering plants.
Despite our significant improvements in
understandiug the patterns and timescale of
angiosperm evolution, the field is far from
settled. The celebrated consistent, strongly
supported phylogeny based on chloroplast
markers is increasingly being recognised as
only one estimate of the angiosperm
phylogeny. The alternative phylogenies
inferred through analysis of nuclear markers,
and through the choice of phylogeny
reconstmction methods, suggests that more
work is needed to reconcile potentially
conflicting evolutionary histories.
Additionally, the controversy surrounding
the age of flowering plants shows no signs of
abating. Modern knowledge of the fossil
record suggests that the rapid radiation of
angiosperm lineages was not quite as
explosive as implied by Darwin’s
“abominable mystery” proclamation, yet a
new mystery is why molecular date estimates
stiU generally far pre-date the oldest
angiosperm fossils. It is unlikely that
increasing the amount of genetic data will
solve this problem (Foster et al. 2016);
instead, increased sampling from
underrepresented groups and
methodological improvements in
incorporating fossil data appear to be the
way forward. The last point appears to be an
especially promising avenue of research, with
new methods being developed for the
simultaneous analysis of extant and extinct
taxa (Ronquist et al. 2012; Gavryushkina et
al. 2014; Heath et al. 2014). Overall, it is
clear that our understanding of the
evolutionary history of angiosperms has
changed considerably over time, and we are
now in an exciting new era of angiosperm
research.
Acknowledgements
I would like to thank Simon Ho for helpful
feedback on this manuscript.
References
Angiosperm Phylogeny Group, A. P. G. (1998)
An ordinal classification for the families of
flowering plants, A.nnals of the Missouri botanical
Garden, 85, 531-553.
Angiosperm Phylogeny Group, A. P. G. (2003)
An update of the Angiosperm Phylogeny
Group classification for the orders and families
of flowering plants. Botanical Journal of the Unnean
Society, 141, 399-436.
Angiosperm Phylogeny Group, A. P, G, (2009)
An update of the Angiosperm Phylogeny
Group classification for the orders and families
of flowering plants: APG III, Botanical Journal of
the Unnean Society, 165, 105—121.
Angiosperm Phylogeny Group, A. P. G. (2016)
An update of the Angiosperm Phylogeny
Group classification for the orders and families
of flowering plants: APG IV, Botanical Journal of
the Unnean Society, 181, 1, 1—20.
Arber, E. A. & Parkin,}. (1907) On the origin of
angiosperms. Journal of the Unnean Society of
Undon, Botany, 38, 263, 29—80.
Beaulieu, J. M., O’Meara, B., Crane, P. &
Donoghue, M. J. (2015) Heterogeneous rates of
molecular evolution and diversification could
explain the Triassic age estimate for
angiosperms, Systematic Biology, 64, 5, 869—878.
Bebber, D. P., Carine, M. A., Wood, J. R. L,
Wortley, A. H., Harris, D. J., Prance, G. T.,
Davidse, G., Paige, J., Pennington, T. D. &
Robson, N. K. B. (2010) Herbaria are a major
frontier for species discovery. Proceedings of the
11
jouRN.\L & Proceedings of the Royal Society of New South Wales
Foster — History of Flowering Plants
National Academy of Sciences^ 107, 51, 22169-
2217L
BeU, C, D., Soltis, D. E. & Soltis, P. S. (2010) The
age and diversification of the angiosperms re¬
revisited, Journal of Botany, 97, 8, 1296—
1303.
BeUard, C., Bertelsmeier, C., Leadley, P., ThuiUer,
W. & Courchamp, F. (2012) Impacts of climate
change on the hiture of biodiversity, 'Ecology
Utters, 15, 4, 365-377.
Bond, W. J. (1989) The tortoise and the hare:
ecology of angiosperm dominance and
g}^mnosperm persistence, Biological Journal of the
Unnean Society, 36, 3, 227-249.
Bowe, L. M., Coat, G. & dePamphilis, C. W.
(2000) Phylogeny of seed plants based on all
three genomic compartments: Extant
gymno sperms are monophyletic and Gnetales'
closest relatives are conifers, Pwceedings of the
National Academy of Sciences, 97, 8, 4092-4097.
Brodribb, T. J. & Feild, T. S. (2010) Leaf
hydraulic evolution led a surge in leaf
photo synthetic capacity during early
angiosperm diversification, Ecology Utters, 13, 2,
175-183.
Bungard, R. A. (2004) Photosynthetic evolution
in parasitic plants: insight from the chloroplast
genome. Bioessays, 26, 3, 235-247.
Burger, W. C. (1977) The Piperales and the
monocots, The Botanical Uview, 43, 3, 345-393.
Burger, W. C. (1981) Heresy revived: the
monocot theory of angiosperm origin.
Evolutionary Theory, 5, 40, 189-226.
Cantino, P. D., Doyle, J. A., Graham, S. W., Judd,
W. S., Olmstead, R. G., Soltis, D. E., Soltis, P. S.
& Donoghue, M. J. (2007) Towards a
phylogenetic nomenclature of Tracheophyta,
Taxon, 56, 3, 1E^4E.
Carmichael, J. S. & Friedman, W. E. (1996)
Double fertilisation in Gnetum gnemon
(Gnetaceae): its bearing on the evolution of
sexual reproduction within the Gnetales and the
anthophyte clade, American Journal of Botany, 83,
6, 767-780.
Chase, M. W., Soltis, D. E., Olmstead, R. G.,
Morgan, D., Les, D. H., Mishler, B. D., Duvall,
M. R, Price, R. A., Hills, H. G. & Qiu, Y.-L.
(1993) Phylogenetics of seed plants: an analysis
of nucleotide sequences from the plastid gene
rAL, Annals of the Missouri Botanical Garden, 80, 3,
528-580.
Chaw, S.-M., Parkinson, C. L., Cheng, Y.,
Vincent, T. M. & Palmer, J. D. (2000) Seed
plant phylogeny inferred from aU three plant
genomes: monophyly of extant gymnosperms
and origin of Gnetales from conifers. Proceedings
of the National Academy of Sciences, 97, 8, 4086-
4091.
Chaw, S.-M., Zharkikh, A., Sung, H.-M., Lau, T.-
C. & Li, W.-H. (1997) Molecular phylogeny of
extant gymnosperms and seed plant evolution:
analysis of nuclear 18S rRNA sequences.
Molecular Biology and Evolution, 14, 1, 56-68.
Crane, P. R. (1985) Phylogenetic analysis of seed
plants and the origin of angiosperms. Annals of
the Missouri Botanical Garden, 72, 4, 716-793.
Cronquist, A. (1981) An integrated system of
classification of flomring plants, Columbia University
Press, New York City, USA.
Darwin, F. & Seward, A. C. (1903) More letters of
Gharles Darwin: a record of his work in a series of
hitherto unpublished letters, John Murray, London,
UK.
Donoghue, M. J. & Doyle, J. A. (1989a)
"Phylogenetic analysis of angiosperms and the
relationships of Hamamelidae", in Crane, P. R.
& Blackmore, S. (eds.) Evolution, systematics, and
fossil history of the Hamamelidae. UoL 1. Introduction
and ”Uwed’ Hamamelidae., Clarendon Press,
Oxford.
Donoghue, M. J. & Doyle, J. A. (1989b)
"Phylogenetic studies of seed plants and
angiosperms based on morphological
characters", in Fernhokn, B., Bremer, K. &
JornvaU, H. (eds.) The hierarchy of life: molecules and
morphology in phylogenetic analysis, Elsevier Science,
Amsterdam, 181-193.
Doyle, J. A. (1969) Cretaceous angiosperm pollen
of the Atlantic Coastal Plain and its
evolutionary significance. Journal of the Arnold
Arboretum, 50, 1—35.
Doyle, J. A. (2012) Molecular and fossil evidence
on the origin of angiosperms. Annual Uview of
Earth and Planetary Sciences, 40, 301-326.
Doyle, J. A. & Donoghue, M. J. (1986) Seed plant
phylogeny and the origin of angiosperms: an
experimental cladistic approach. The Botanical
Review, 52, 4, 321-M-3L
78
Journal & Proceedings of the Royal Society of New South Wales
Foster — History of Flowering Plants
Doyle, J. A. & Hotton, C. L. (1991)
"Diversification of early angiosperm poUen in a
cladistic context", in Blackmore, S. & Barnes, S.
H. (eds.) 'Pollen and spores: patterns of diversification^
Clarendon Press, Oxford, 169-195.
Ellis, R. J. (2010) Biochemistry: Tackling
unintelligent design, Nature, 463, 7278, 164-165.
Endress, P. K. (1987) The early evolution of the
angiosperm flower. Trends in Ecologji Evolution,
2, 10, 300-304.
Endress, P. K. (2002) Morphology and
angiosperm systematics in the molecular era.
The Botanical Rfvieiv, 68, 4, 545-570.
Endress, P. K., Baas, P. & Gregory, M. (2000)
Systematic plant morphology and anatomy: 50
Years of progress. Taxon, 49, 3, 401-434.
Fakon-Demaret, M. (1996) Dorinnotheca streelii
Fairon-Demaret, et sp. nov., a new early seed
plant from the upper Famennian of Belgium,
Review of Palaeobotanj and Paljnok^, 93, 1, 217-
233.
Fairon-Demaret, M. & Scheckler, S. E. (1987)
Typification and redescription of Moresnetia
c(alesskji Stockmans, 1948, an early seed plant
from the Upper Famennian of Belgium, Bulletin-
Institut Royal des Sciences Naturelles de Belgique.
Sciences de la Terre, 57, 183-199.
Fleischmann, A., Michael, T. P., Rivadavia, F.,
Sousa, A., Wang, W., Temsch, E. M.,
Greilhuber, J., MiiUer, K. F. & Heubl, G. (2014)
Evolution of genome size and chromosome
number in the carnivorous plant genus Genlisea
(Lentibulariaceae), with a new estimate of the
minimum genome size in angiosperms, Annals
ofBotany, 114, 8, 1651-1663.
Foster, C. S. P., Sauquet, H., van der Merwe, M.,
McPherson, H., Rossetto, M. & Ho, S. Y. W.
(2016) Evaluating the Impact of Genomic Data
and Priors on Bayesian Estimates of the
Angiosperm Evolutionary Timescale, Systematic
Biolog)!, in press.
Friedman, W. E. (1992) Evidence of a pre-
angiosperm origin of endosperm: implications
for the evolution of flowering plants, Science,
255, 5042, 336-339.
Galtier, N. (2011) The intriguing evolutionary
dynamics of plant mitochondrial DNA, BMC
Biolog)!, 9, 1, 61.
Gang, H. A. N., Zhongjian, L. 1. U., Xueling, L. 1.
U., Limi, M. A. O., Jacques, F. & Xin, W.
(2016) A Whole Plant Herbaceous Angiosperm
from the Middle Jurassic of China, Acta
Geologica Sinica (English Edition), 90, 1, 19-29.
Gavryushkina, A., Welch, D., Stacker, T. &
Dmmmond, A. J. (2014) Bayesian inference of
sampled ancestor trees for epidemiology and
fossil calibration, P1j)S Computational Biology, 10,
12, el003919.
Goremykin, V. V., Hansmann, S. & Martin, W. F.
(1997) Evolutionary analysis of 58 proteins
encoded in six completely sequenced
chloroplast genomes: revised molecular
estimates of two seed plant divergence times.
Plant Systematics and Evolution, 206, 1—4, 337—351.
Goremykin, V. V., Nikiforova, S. V., Biggs, P. J.,
Zhong, B., Delange, P., Martin, W., Woetzel, S.,
Atherton, R. A., McLenachan, P. A. &
Lockhart, P. J. (2013) The evolutionary root of
flowering plants, Systematic Biology, 62, 1, 50—61.
Goremykin, V. V., Nikiforova, S. V., Cavalieri,
D., Pindo, M. & Lockhart, P. (2015) The Root
of Flowering Plants and Total Evidence,
Systematic Biolog!, 64, 5, 879-891.
Govaerts, R. (2001) How many species of seed
plants are there?, Taxon, 50, 4, 1085-1090.
Hay, W. W. & Floegel, S. (2012) New thoughts
about the Cretaceous climate and oceans, Earth-
Science Reviews, 1 15, 4, 262-272.
Heath, T. A., Huelsenbeck, J. P. & Stacker, T.
(2014) The fossilized birth-death process for
coherent calibration of ckvergence-time
estimates. Proceedings of the National Academy of
Sciences, 111, 29, E2957-E2966.
Ho, S. Y. W. (2014) The changing face of the
molecular evolutionary clock. Trends in Ecology
(Er Evolution, 29, 9, 496-503.
Ho, S. Y. W. & Duchene, S. (2014) Molecular-
clock methods for estimating evolutionary rates
and Molecular Ecology, 23, 5947-5965.
Ho, S. Y. W. & Phikips, M. J. (2009) Accounting
for cakbration uncertainty in phylogenetic
estimation of evolutionary ckvergence times,
Systematic Biolog, 58, 3, 367-380.
Ho, S. Y. W., Tong, K. J., Foster, C. S. P., Ritchie,
A. M., Lo, N. & Crisp, M. D. (2015)
Biogeographic calibrations for the molecular
clock, Biolog Letters, 11,9, 20150194.
Hochuk, P. A. & Feist-Burkhardt, S. (2013)
Ang^osperm-kke poken and Afropollis from the
Midcke Triassic (Anisian) of the Germanic
79
jouRN.\L & Proceedings of the Royal Society of New South Wales
Foster ~ History of Flowering Plants
Basia (Northern Switzerland), Frontiers in Plant
Science^ 4, 344.
Jiao, Y., Wickett, N. J., Ayyampalayam, S.,
Chanderbali, A. S., Landherr, L., Ralph, P. E.,
Tomsho, L. P., Hu, Y., Liang, H. & Soltis, P. S.
(2011) Ancestral polyploidy in seed plants and
angiosperms. Nature, 473, 7345, 97-100.
Kitazaki, K. & Kubo, T. (2010) Cost of having
the largest mitochondrial genome: evolutionary
mechanism of plant mitochondrial genome,
Journal of Botanj, 20 1 0, 620 1 3 7 .
Kjrassilov, V. A. (1977) The origin of
angiosperms. The Botanical Review, 43, 1, 143—
176.
Lanfear, R, Ho, S. Y. W., Davies, T. J., Moles, A.
T., Aarssen, L., Swenson, N. G., Warman, L.,
Zanne, A. E. & AUen, A. P. (2013) Taller plants
have lower rates of molecular evolution, Nature
Communications, 4, 1879.
Laroche, J., Li, P. & Bousquet, J. (1995)
Mitochondrial DNA and monocot-dicot
divergence time. Molecular Biolo^ and Evolution,
12, 6,1151-1156.
Lee, M. S. Y. & Ho, S. Y. W. (2016) Molecular
clocks. Current Biology, 26, 10, R399-R402.
Levin, D. A. (2002) The role of chromosomal change in
plant evolution, Oxford University Press, New
York.
MagaUon, S. (2014) A review of the effect of
relaxed clock method, long branches, genes,
and calibrations in the estimation of angiosperm
age. Botanical Sciences, 92, 1, 1-22.
MagaUon, S. & Sanderson, M. J. (2001) Absolute
diversification rates in angiosperm clades.
Evolution, 55, 9, 1762-1780.
MagaUon, S., Gomez-Acevedo, S., Sanchez-
Reyes, L. L. & Hernandez-Hernandez, T.
(2015) A metacaUbrated time- tree documents
the early rise of flowering plant phylogenetic
diversity, New Phjtologist, 207, 2, 437-453.
Martin, W., Gierl, A. & Saedler, H. (1989)
Molecular evidence for pre-Cretaceous
angiosperm origins. Nature, 339, 46-48.
Mathews, S. & Donoghue, M. J. (1999) The root
of angiosperm phylogeny inferred from
dupUcate phytochrome genes. Science, 286, 5441,
947-950.
Meeuse, A. D. J. (1972) Facts and fiction in floral
morphology with special reference to the
Polycarpicae 1. A general survey, Acta Botanica
Neerlandica, 21, 2, 113-127.
Moore, M. J., BeU, C. D., Soltis, P. S. & Soltis, D.
E. (2007) Using plastid genome-scale data to
resolve enigmatic relationships among basal
angiosperms. Proceedings of the National Academy of
Sciences, 104, 49, 19363-19368.
Moore, M. J., Soltis, P. S., BeU, C. D., Burleigh, J.
G. & Soltis, D. E. (2010) Phylogenetic analysis
of 83 plastid genes further resolves the early
diversification of eudicots, Proceedings of the
National Academy of Sciences, 107, 10, 4623-4628.
Parkinson, C. L., Adams, K. L. & Palmer, J. D.
(1999) Multigene analyses identify the three
earUest lineages of extant flowering plants.
Current Biologf, 9, 24, 1485-1491.
PeUicer, J., Fay, M. F. & Leitch, 1. J. (2010) The
largest eukaryotic genome of them aU?, Botanical
Journal of the Einnean Society, 1 64, 1 , 1 0—1 5.
Pimm, S. L. & Joppa, L. N. (2015) How many
plant species are there, where are they, and at
what rate are they going extinct?. Annals of the
Missouri Botanical Garden, 100, 3, 170—176.
Preston, J. C. & HUeman, L. C. (2009)
Developmental genetics of floral symmetry
evolution. Trends in Plant Science, 14, 3, 147-154.
Qiu, Y.-L., Lee, J., Bernasconi-Quadroni, F.,
Soltis, D. E., Soltis, P. S., Zanis, M., Zimmer, E.
A., Chen, Z., Savolainen, V. & Chase, M. W.
(1999) The earUest angiosperms; evidence from
mitochondrial, plastid and nuclear genomes.
Nature, 402, 6760, 404-407.
Ray, J. (1686-1704) Historia Plantarum, H.
Faithorne, London.
Ronquist, F., Klopfstein, S., Vilheknsen, L.,
SchuUneister, S., Murray, D. L. & Rasnitsyn, A.
P. (2012) A total-evidence approach to dating
with fossUs, appUed to the early radiation of the
Hymenoptera, Systematic Biology, 61, 6, 973-999.
Rosas-Guerrero, V., AguUar, R., Marten-
Rodriguez, S., Ashworth, L., Lopezaraiza-Mikel,
M., Bastida, J. M. & Quesada, M. (2014) A
quantitative review of pollination syndromes:
do floral traits predict effective pollinators?.
Ecology Fetters, 17, 3, 388-400.
RothweU, G. W., Scheckler, S. E. & GiUespie, W.
H. (1989) Elkinsia gen. nov., a Late Devonian
gymnosperm with cupulate ovules, Botanical
Gazette, 150, 2, 170-189.
80
Journal & Proceedings of the Royal Society of New South Wales
Foster — History of Flowering Plants
Ruhfel, B. R., Gitzendanner, M. A., Soltis, P. S.,
Soltis, D. E. & Burleigh, J. G. (2014) From
algae to angiosperms—mferring the phylogeny
of green plants (Viridiplantae) from 360 plastid
genomes, BMC Evolutionary Biology, 14, 1, 23.
Saarela, J. M., Rai, H. S., Doyle, J. A., Endress, P.
K., Mathews, S., Marchant, A. D., Briggs, B. G.
& Graham, S. W. (2007) HydateUaceae
identified as a new branch near the base of the
angiosperm phylogenetic tree. Nature, 446,
7133,312-315.
Sagan, L. (1967) On the origin of mitosing ceUs,
Journal of Theoretical Biology, 14, 3, 225-274.
Sage, R. F., Christin, P.-A. & Edwards, E. J.
(2011) The C4 plant lineages of planet Earth,
Journal of Experimental Botanj, 62, 9, 3155-3169.
Sanderson, M. J. & Doyle, J. A. (2001) Sources of
error and confidence intervals in estimating the
age of angiosperms from rAL and 18S rDNA
data, American Journal of Botany, 88, 8, 1499—
1516.
Schwartz, R. M. & Dayhoff, M. O. (1978) Origins
of prokaryotes, eukaryotes, mitochondria, and
chloroplasts, 199, 4327, 395^03.
Scotland, R. W. & Wordey, A. H. (2003) How
many species of seed plants are there?. Taxon,
52, 1, 101-104.
Smith, S. A., Beaulieu, J. M. & Donoghue, M. J.
(2010) An uncorrelated relaxed-clock analysis
suggests an earlier origin for flowering plants.
Proceedings of the National Academy of Sciences, 107,
13, 5897-5902.
Smith, S. A. & Donoghue, M. J. (2008) Rates of
molecular evolution are linked to life history in
flowering plants. Science, 322, 5898, 86-89.
Soltis, D. E., Smith, S. A., CeRinese, N., Wurdack,
K. J., Tank, D. C., Brockington, S. F., RefuMo-
Rodriguez, N. F., Walker,}. B., Moore, M. J. &
Carlsward, B. S. (2011) Angiosperm phylogeny:
17 genes, 640 X2cm, American Journal of Botany, 98,
4, 704-730.
Soltis, P. S. & Soltis, D. E. (2004) The origin and
diversification of angiosperms, American Journal
of Botany, 91, 10, 1614—1626.
Soltis, P. S., Soltis, D. E. & Chase, M. W. (1999)
Angiosperm phylogeny inferred from multiple
genes as a tool for comparative biology. Nature,
402, 6760, 402-404.
Soltis, P. S., Soltis, D. E., Savolainen, V., Crane, P.
R. & Barraclough, T. G. (2002) Rate
heterogeneity among lineages of tracheophytes:
integration of molecular and fossil data and
evidence for molecular Hving fossRs, Proceedings
of the National Academy of Sciences, 99, 7, 4430—
4435.
Takhtajan, A. L. (1980) Outiine of the
classification of flowering plants
(MagnoHophyta), The Botanical Review, 46, 3,
225-359.
ThuRler, W., Lavergne, S., Roquet, C.,
Boulangeat, L, Lafourcade, B. & Araujo, M. B.
(2011) Consequences of cRmate change on the
tree of Rfe in Europe, Nature, 470, 7335, 531-
534.
Turcotte, M. M., Davies, T. J., Thomsen, C. J. M.
& Johnson, M. T. J. (2014) Macroecological and
macroevolutionary patterns of leaf herbivory
across vascular plants, Proceedings of the Rnyal
Society of Eondon B: Biological Sciences, 281, 1787,
20140555.
van der Niet, T. «& Jolinson, S. D. (2012)
Phylogenetic evidence for poRinator-driven
diversification of angiosperms. Trends in Ecology
Cn Evolution, 27, 6, 353-361.
Wang, X., Duan, S., Geng, B., Cui, J. & Yang, Y.
(2007) Is Jurassic Schmeissneria an angiosperm?,
Acta Palaeontologica Sinica, 46, 4, 490.
Welch, J. J. & Bromham, L. (2005) Molecular
dating when rates vary, Trends in Ecology
Evolution, 20, 6, 320-327.
Wettstein, R. (1907) Handbuch der systematischen
Botanik, 2 edn., Franz Deuticke, Leipzig.
Wickett, N. J., Mirarab, S., Nguyen, N., Warnow,
T., Carpenter, E., Matasci, N., Ayyampalayam,
S., Barker, M. S., Burleigh, J. G. &
Gitzendanner, M. A. (2014)
Phylotranscriptomic analysis of the origin and
early diversification of land plants, Proceedings of
the National Academy of Sciences, 111, 45, E4859—
E4868.
Winter, K.-U., Becker, A., Munster, T., Kim, J. T.,
Saedler, H. & Theissen, G. (1999) MADS-box
genes reveal that gnetophytes are more closely
related to conifers than to flowering plants.
Proceedings of the National Academy of Sciences, 96,
13, 7342-7347.
Wolfe, K. H., Li, W.-H. & Sharp, P. M. (1987)
Rates of nucleotide substitution vary greatly
among plant mitochondrial, chloroplast, and
81
Journal & Proceedings of the Royal Society of New South Wales
Foster — History of Flowering Plants
nuclear DNAs, Proceedings of the 'National Academy
of Sciences, 84, 24, 9054—9058.
Xi, Z., Liu, L., Rest, J. S. & Davis, C. C. (2014)
Coalescent versus concatenation methods and
the placement of AmboreUa as sister to water
lilies. Systematic Bio logjj, 63, 6, 919-932.
Zanne, A. E., Tank, D. C., CornweU, W. K.,
Eastman, J. M., Smith, S. A., Fitzjohn, R. G.,
McGlinn, D. ]., O’Meara, B. C., Moles, A. T.,
Reich, P. B., Doyer, D. L., Soltis, D., Stevens,
P., Westoby, M., Wright, L, Aarssen, L., Benin,
R., Calaminus, A., Govaerts, R, Hemmings, F.,
Leishman, J., Soltis, P. S., Swenson, N. G.,
Warman, L. & Beaulieu,}. M. (2014) Three keys
to die radiation of angiosperms into freezing
environments. Nature, 506, 7486, 89-92.
Zeng, L., Zhang, Q., Sun, R,, Kong, H., Zhang,
N. & Ma, H. (2014) Resolution of deep
angiosperm phylogeny using conserved nuclear
genes and estimates of early divergence times,
Nature Communications, 5, 4956.
Zhong, B., Yonezawa, T., Zhong, Y. &
Hasegawa, M. (2010) The position of Gnetales
among seed plants: overcoming pitfaUs of
chloroplast phylogenomics. Molecular Biolog)i and
'Evolution, 21, 12, 2855-2863.
Zuckerkandl, E. & Pauling, L. (1962) "Molecular
disease, evolution and genic heterogeneity.", in
Kasha, M. & Pullman, B. (eds.) Horizons in
Biochemistry., Academic Press., New York, 189—
225.
82
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
p. 83. ISSN 0035-9173/16/010083-01
Tliesis abstract
Adherence to self-management and psychological distress in
women with breast cancer-related lymphoedema
Jessica Alcorso
Abstract of a thesis for a Doctorate of Philosophy submitted to Macquarie University, Sydney, Australia
Nonadherence to breast cancer-related
lymphoedema self-management is sub-
optimal. In addition, women with breast
cancer-related lymphoedema experience
psychological distress associated with this
chronic illness. Adopting a social-cogni¬
tive theoretical framework, the aim of this
thesis is to identify cognitive and affective
predictors of adherence to self-management
behaviours and predictors of psychological
distress in women living with breast cancer-
related lymphoedema. To achieve this aim,
a longitudinal study was conducted that
measured adherence to self-management
behaviours, psychological distress, and
cognitive and affective factors at baseline,
6- and 12-months. An additional cross-
sectional study was conducted to identify
lymphoedema therapists’ and affected
women’s beliefs about barriers to adherence
to self-management. The findings from
the empirical studies on adherence suggest
that cognitive and affective factors are not
informative for understanding self-man¬
agement behaviour in women with breast
cancer-related lymphoedema. Furthermore,
a disconnect between therapists’ and affec¬
tive women’s beliefs about barriers to self¬
management was identified, with therapists
believing more strongly than the affected
women that financial cost, time, concerns
about appearance, difficulty accessing treat¬
ment, insufficient knowledge, and physical
limitations negatively impact adherence. In
contrast, a number of cognitive and affective
factors significantly predicted distress. Based
on the results from the longitudinal study, an
online self-compassion based writing activ¬
ity was developed to minimise distress and
body image disturbance in this population of
lymphoedema-affected women. The online
intervention received moderate to high user
acceptability ratings from women affected
with breast cancer-related lymphoedema
suggesting the potential viability of this
intervention. In sum, the findings from
this thesis have important implications for
researchers and health professionals. Regard¬
ing self-management, medical characteristics
and knowledge were identified as important
factors for identifying women at risk of non¬
adherence. In addition, it may be beneficial
to screen women for symptoms of psycho¬
logical distress and body image disturbance
in order to identify who may benefit from
additional psychosocial support.
Jessica Alcorso,
Department of Psychology,
Macquarie University,
Sydney NSW 2109AUSTRALIA
Email: jessica.alorso@mq.edu.au
83
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
p. 84. ISSN 0035-9173/16/010084-01
Thesis abstract
A social psychological examination of factors shaping
career and education aspirations through childhood and
adolescence
Nathan Berger
Abstract of a thesis for a Doctorate of Philosophy submitted to University of Newcastle, Newcastle,
Australia
Young people often are asked what they
want to be when they grow up. What fac™
tors influence their responses to this most
common of questions? This study used
theoretical perspectives from social and
vocational psychology, including Gottfred-
sons (1981, 2002) career circumscription
and compromise theory, to investigate the
formation of career aspirations during pri¬
mary and secondary schooling. A particu¬
lar emphasis was the relationship between
social cognition and socioeconomic status
(SES) is SES associated with young peo¬
ple s thinking about careers-related identity
questions? The study drew on data gathered
during the first three years (2012-2014) of
the Aspirations Longitudinal Study (ALS)
in the State of New South Wales, Australia.
Students in Years 3-11 were sampled in an
accelerated longitudinal design, whereby
three cohorts completed a comprehensive
career aspirations questionnaire for three
years. The sample comprised 7,462 ques¬
tionnaires from 5j304 students. In addi¬
tion to the questionnaires, the ALS accessed
standardised demographic and literacy/
numeracy data held by the state for each
student. To examine the influence of stu¬
dents’ social-cognitive styles on their career
aspirations, they also completed a modified
version of Berzonsky et ah s (2013) Identity
Style Index as part of the 2014 questionnaire.
The latent class mixture modelling (LCMM)
technique was used to analyse the acceler¬
ated longitudinal data. LCMM identified
four discrete change trajectories in the pres¬
tige of career aspirations during nine years
of schooling. Students’ identity styles were
found to differ between SES backgrounds,
with high SES students more information-
oriented and less diffiise/ avoidant when deal¬
ing with identity questions compared to low
SES students. Identity styles also differed by
SES background within the LCMM trajec¬
tories, with high aspiring low SES students
more information-oriented and less diffuse/
avoidant than their low aspiring peers. The
implications of these findings for theory
research, and practice are discussed.
Dr Nathan Berger,
School of Education,
University of Newcastle,
Newcastle NSW 2300
AUSTRALIA
Email: n.berger@westernsydney.edu.au
84
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 20 1 6,
pp. 85-86. ISSN 0035-9173/16/010085-02
Thesis abstract
The cognitive and neural mechanisms of joint attention:
a second person approach
Nathan Caruana
Abstract of a thesis for a Doctorate of Philosophy submitted to Macquarie University, Sydney, Australia
This project investigated the cognitive and
neural mechanisms of joint attention in typi¬
cal development and in high-functioning
autism. A novel gaze-contingent virtual real¬
ity tasks was developed and implemented
in a number of studies using a range of
techniques, including functional magnetic
resonance imaging (fMRI), event-related
potentials (ERP) and eye tracking.
The first aim of this project was to develop
an experimental joint attention paradigm
that (1) captured both the initiating and
responding functions of joint attention, (2)
could be applied in both behavioural and
neurophysiological experiments, (3) pro¬
vided full experimental control over non¬
social task demands, and (4) established an
ecologically valid context for joint attention
interactions.
Chapter 2 of this dissertation reviewed
the current approaches for measuring joint
attention in experimental settings and intro¬
duced a new virtual reality paradigm of joint
attention that achieves both experimental
control and ecological validity. Critical
issues associated with the measurement of
joint attention are discussed.
The second aim of this project was to use
this paradigm to investigate the neural and
cognitive mechanisms that support joint
attention in typical development and in
autism.
In Chapter 3, fMRI was used to investi¬
gate the neural correlates that were unique
and common to initiating and responding
to joint attention bids in 13 adults with typi¬
cal development. A right-lateralised fronto-
temporoparietal network was found to be
common to both initiating and respond¬
ing to joint attention bids and comprised
the middle frontal gyrus (MFC), inferior
frontal gyrus (IFG), middle temporal gyrus
(MTG), precentral gyrus, posterior superior
temporal sulcus (pSTS), temporoparietal
junction (TPJ) and precuneus. Compared
to responding to joint attention bids, ini¬
tiating joint attention was associated with
additional activation of the MFG, IFG, TPJ
and precuneus.
In Chapter 4, eye-tracking was used to
investigate joint attention performance in 17
adults with high-functioning autism and 1 7
adults with typical development (controls).
Individuals with autism made significantly
more errors than controls when initiating
and responding to joint attention bids.
In Chapter 5, another virtual reality para¬
digm was developed and employed in an
event-related potential study which inves¬
tigated the time course of neural processes
associated with evaluating self-initiated joint
85
Journal & Proceedings of the Royal Society of New South Wales
Camana — Mechanisms of Joint Attention
attention bids. In a sample of 19 partici¬
pants with typical development, centro-pari-
etal P350 and P500 ERPs were significantly
larger when gaze shifts resulted in the avoid¬
ance, rather than the achievement of joint
attention. This P350 and P500 morphology
was absent in a second sample of 1 9 partici¬
pants who completed a non-social version of
the same task in which arrows replaced the
gaze of the virtual character.
In Chapter 6, the same paradigm was
used to investigate whether the P350 effect
observed in Chapter 5 was influenced by
participants’ beliefs of their virtual partner s
agency. The P350 effect was only observed
in participants who believed that their part¬
ner was controlled by a human (n - 19), and
not in a second group of individuals who
were informed that the virtual character
was controlled by a computer program (n
= 19).
Nathan Caruana,
Department of Cognitive Science,
Macquarie University,
Sydney NSW 2109
Australia
Email: Nathan.caruana@mq.edu.au
86
Journal dr Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
p. 87. ISSN 0035-9173/16/010087-01
Thesis abstract
Environmental flows at work; restoring floodplain wedands
through return of historical conditions
Samantha I^therine Dawson
Abstract of a thesis for a Doctorate of Philosophy submitted to the University of New South Wales,
Sydney, Australia
Wetlands are among the most degraded
ecosystems worldwide, demanding effec¬
tive restoration. In an effort to ameliorate
ecological degradation from upstream water
diversions, environmental and managed
flows are increasingly used to help restore
vegetation communities. Understanding
of factors affecting the success and efficacy,
however, remains limited and is of increasing
urgency as this type of restoration becomes
more widespread. I investigated the capacity
of flooding, including environmental flows,
to restore wetland flora across areas varying
in degradation from historic land-use.
I predicted that increasing land-use his¬
tory (increasing duration and decreasing
time since land use) would decrease resto¬
ration effectiveness, while increased flood¬
ing frequency and duration should improve
restoration outcomes. A case study was used
to assess vegetation restoration: a floodplain
wetland with fields representing a land-use
chronosequence, flooded through environ¬
mental flows, in the Macquarie Marshes in
the Murray-Darling Basin of south-eastern
Australia. I examined extant vegetation, soil
seedbanks, plant trait distributions and his¬
torical vegetation change through surveys of
plant community composition, greenhouse
seed bank germination, fourth-corner trait
modelling approaches and Landsat imagery
analysis.
Composition of extant herbaceous veg¬
etation correlated with both land-use history
and flooding, while extant shrub and woody
species were more strongly correlated with
land-use. Within seedbanks, exotic and dis¬
turbance-adapted species were associated with
increased duration and decreased time since
land-use and native wetland species were asso¬
ciated with opposite land-use practice and
increased flooding. Furthermore, I found that
there was capacity within the soil seed bank
to engender further restoration of extant veg¬
etation. Trait analyses showed native, woody
and clonal species were taking the longest to
restore, especially in high land use areas, but
this may be ameliorated through increased
inundation. Landsat analyses demonstrated
that inundation was vital to restoration and
also indicated a gradient of restoration success,
with areas of less land use history (e.g. clearing
and one year of cultivation) restoring more
quickly than longer cultivation durations. In
conclusion, flooding was critical to achieving
restoration objectives, with higher frequen¬
cies increasing efficacy, but increased land-use
intensity compromises restoration rates and
possibly success.
Dr Samantha Dawson,
Centre for Ecosystem Science,
University of New South Wales,
Sydney NSW 2052 Australia
Email: samantha.k.dawson@gmail.com
Thesis: handle.unsw.edu.au/ 1 959.4/ 56248
87
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pp. 88-89. ISSN 0035-9173/16/010088-02
Thesis abstract
Quantitative proteomic analyses of isolate variation and
virulence in Giardia duodenalis
Samantha J Emety
Abstract of a thesis for a Doctorate of Philosophy submitted to Macquarie University, Sydney, Australia
Giardia duodenalis is a parasitic protozoan
with a global human infection burden of
250 million, and is therefore the largest para¬
sitic cause of diarrheal disease worldwide.
Though some cases are asymptomatic, giar¬
diasis can be acute and chronic, with post¬
infection sequellae including irritable bowel
syndrome, chronic fatigue, obesity and type
II diabetes. Importantly, Giardia is problem¬
atic in children under the age of five, causing
ill-thrift and failure-to-thrive. In addition,
diarrheal diseases including Giardia consti¬
tute the second-leading cause of mortality
for this age category Giardia has a direct life
cycle, where infective, tetranucleated cysts
are transmitted via the faeco-oral route, and
then excyst in the duodenum into virulent,
flagellated trophozoites. The prevalence of
the parasite is also due to its wide host range,
with zoonotic transfer from wild, livestock
and domestic animal species to humans.
Efforts continue to define the mechanisms
of virulence and pathophysiology, as more
research is needed to elucidate the relation¬
ship between host and parasite factors.
Advances in genetic epidemiology have
defined clear assemblages that segregate
phylogenetically according to host range,
and multiple assemblage and subassem¬
blage genome sequences are now available.
These genome sequences have provided the
databases necessary for bottom-up, or shot¬
gun, proteomics, and as such have expanded
possibilities for quantitative analyses in this
parasite. This thesis aimed to provide a
thorough quantitative proteomic founda¬
tion to enhance the Giardia research field
both biologically and technically. To achieve
this, the thesis consists of four experimental
investigations into aspects of parasite varia¬
tion and virulence, all of which have gener¬
ated quantitative proteomic data.
Firstly, two different protein sample
preparation and fractionation methods
were compared for label-free quantitative
proteomics. Tliese were applied to two G.
duodenalis assemblage A1 isolates with dif¬
ferent phenotypes, in order to investigate
possible sources of isolate variation. The
optimised protocol generated from this ini¬
tial investigation was applied in later studies,
which are also contained within this thesis.
In addition, phenotypes associated with
pathogenicity correlated with up-regulation
of known virulence factors in Giardia.
Following this initial investigation, quanti¬
tative data was generated using the same label-
free approach for eight assemblage A isolates,
which constituted the first comprehensive
proteomic baseline for this taxonomic group.
Isolates of diverse host, geographic and sub¬
assemblage origins were analysed using mass
spectrometry to characterise their common
proteomes and isolate-specific variations. In
addition, both the A1 and A2 subassemblage
genome databases were evaluated for peptide
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Journal & Proceedings of the Royal Society of New South W^es
Emery— “Variation and Virulence in Giardia duodenalis
to spectrum matching, which demonstrated
the importance of subassemblage databases
to improve identifications from the Giardia
variable genome.
The third study investigated isolate varia¬
tion in the biological context of the process
of differentiation in G. duodenalis. Label-
free quantitative proteomics was used to ana¬
lyse the proteomes of cysts and trophozoites
from two genome-alternate subassemblage
A1 isolates. This is the first post-genomic
analysis of the life cycle beyond the genome
isolate, WB. A range of isolate-independent,
universal encystation markers were identi¬
fied, as well as several indications of isolate-
specific life-cycle adaptations which may
impact reinfection success in subsequent
generations.
Finally, the last experiment in this thesis
investigated disease induction using in vitro
host-parasite interaction models between
intestinal epithelial cell (lEC) lines and
trophozoites. We used isobaric Tandem
Mass Tags (TMT) to sensitively quantitate
changes in trophozoites which were either
allowed to attach to host-cell monolayers,
or were exposed to host-cell secretions alone.
This is the first use of TMT label technolo¬
gies for quantitative proteomics in Giardia.
This has demonstrated that distinct protein
cascades are induced by both levels of host-
signals, and also that induction of virulence
factors is not dependent on parasite attach¬
ment to host cells.
Through these experiments, this thesis
demonstrates that a range of quantitative
proteomic approaches are suitable for G.
duodenalis, all of which are capable of pro¬
viding important insight into key aspects
of parasite biology. These studies provide
an important proteomic complement for
genomic and transcriptomic data currently
available in the literature, which is necessary
for undertaking a systems biology approach
to understanding Giardia.
Dr Samantha J Emery,
Department of Chemistry and Biomolecular
Sciences,
Macquarie University,
Sydney NSW 2109
AUSTRALIA
Email: emery.s(2)wehi. edu.au
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Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 90-91. ISSN 0035-9173/16/010090-02
Thesis abstract
Improving labour outcomes in the creative industries:
the role of creative workers’ social network structure and
organisational business acumen
Benjamin Farr-Wharton
Abstract of a thesis for a Doctorate of Philosophy submitted to Southern Cross University, Lismore,
Australia
Individuals who work in artistic, cultural
and creative fields (henceforth creative
workers) are increasingly conceptualised as
a labour force. Under this conceptualisa¬
tion, creative workers are seen to contribute
positively to significant, national economic
indicators such as Gross Domestic Product
and innovation indices. However, a grow¬
ing body of research indicates that, as a
labour force, ‘creative workers’ experience
particularly poor labour outcomes^ insofar as
they typically work longer hours, for rela¬
tively lower incomes, and have very little
job security.
The theoretical concepts provided by the
Resource-Based View (RBV) of the firm and
Bourdieu’s theories concerning social and
cultural capital, and the field of cultural pro¬
duction, are used in this thesis to examine
the impact of organisational business acumen
and social network structure on the labour
outcomes (exploitation, labour precarity and
earnings satisfaction) of creative workers. The
research adopts a mixed methods approach
that encompasses an initial pilot case study
using social network analysis, followed by an
explanatory quantitative-qualitative design.
The study is situated within an Australian
context, and a total of three hundred people
were involved in the data collection for the
sum of these three research phases.
Results from the quantitative and quali¬
tative research indicate that organisational
business acumen and social network structure
significantly reduce the labour precarity of
creative workers. In addition, organisational
business acumen significantly reduces per¬
ceptions of exploitation, as well as enhances
the earnings satisfaction of sampled creative
workers.
This research is innovative and impor¬
tant in a number of ways. For practitioners,
results from this thesis highlight the signifi¬
cance of both social network structure and
organisational business acumen as resources
that positively impact the labour outcomes
of creative workers. For policy makers,
the research encourages the development
of new policy instruments to support the
diffusion of organisational business acumen
across the creative labour force, in order
to enhance the efficiency of the creative
sector. For theory, the research suggests
that creative workers are no longer subject
to a dichotomous arts’ or ‘market’ mode of
production, but compromise business prac¬
tices and artistic pursuit to seek sustainable
outcomes. For the field of network analysis,
the research offers new quantitative instru-
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Journal & Proceedings of the Royal Society of New South Wales
Farr-Wharton— Labour Outcomes in the Creative Industries
ments (specifically catering for the context
of the network-centric creative sector) to
measure the impact of network structures
on performance measures.
Dr Benjamin Farr-Wharton
School of Business and Tourism
Southern Cross University
Lismore NSW 2480
AUSTRALIA
Email: 128154@uts.edu.au
91
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 92-93. ISSN 0035-9173/16/010092-02
Thesis abstract
Dim and dimmer: the production and difhision
of the natural sciences in Australia
between the 1770s and the 2010s
Lynnette Hicks
Abstract of a thesis for a Doctorate of Philosophy submitted to Macquarie University, Sydney, Australia
Despite growing public concerns around
socio-scientific problems and the significance
of these problems to everyday life, there is a
dearth of sociological literature addressing
the production and diffusion of the natural
sciences in Australia. In particular, critical
analyses of scientific knowledge production
and diffusion relative to the actions of the
state, the market and civil society are largely
absent. This thesis sets out to mitigate this
situation by contributing a critical histori¬
ography of scientific knowledge production
and diffusion as it relates to Australia since
white settlement. It is anticipated that this
work will open up the topic for further aca¬
demic research and rational debate.
This thesis explores the production and
diffusion of scientific knowledge through the
lens of social dynamics that have emerged in
Australia between the 1770s and the 2010s.
The research relies primarily on the theoreti¬
cal work of Max Weber in order to identify
and analyse the conception of rationality
and its application to social action that is
present in the policy and praxis of the natu¬
ral sciences in Australia. In particular the
relationships between the state, the market
and civil society are analysed using secondary
data drawn from published histories, offi¬
cial documents and the formal policies and
practices of the state and the market during
this period.
A tripartite analytical model has been cre¬
ated specifically for this thesis and is utilised
to trace scientific knowledge production and
diffusion through the transformative social
processes associated with instrumentalism ^
bureaucratisation, developmentalism, envi¬
ronmentalism^ postmodernism and neoliber¬
alism. Rationality is applied in three ways: as
non-instrumental science produced to further
human understandings of the natural world
and to promote the development of civil
society; as pre-instrumental science produced
by the state to in order to develop markets
and for other instrumental purposes such as
national defence strategies; and as instrumen¬
tal scientific knowledge produced by the par¬
ticipants in the market expressly to enhance
their own position in the market.
Tfie research reveals that instrumental
rationality has been an enduring concept in
the policy and praxis of the natural sciences
in Australia. Moreover, this thesis finds that a
strong tension is often present between non¬
instrumental notions of scientific knowledge
and those practices that are predominantly
instrumental. Through each of the periods
studied the state and the market have been
close confederates, often working together
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Journal & Proceedings of the Royal Society of New South Wales
Hicks— The Natural Sciences in Australia
to realize instrumental outcomes through
the knowledge produced by natural science.
In particular, administrative and economic
ends are seen to be primary; ends associated
with more normative intentions, such as the
nurturing of civil society, have been regularly
overlooked in favour of strictly instrumental
aspirations. This continuing instrumental¬
ity has altered the relationships between the
state, the market and civil society during
each period studied. On the current tra¬
jectory, the policy and praxis of the natural
sciences in Australia may yet begin to com¬
promise the sovereignty of that nation state
and the authority of its citizenry.
Dr Lynnette Hicks,
Department of Sociology,
Macquarie University,
Sydney NSW 2109
AUSTRALIA
Email: lyn.hicks@mq.edu.au
lyn.hicks@bigpond.com
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p. 94. ISSN 0035-9173/16/010094-01
Thesis abstract
Pale Communion: whiteness, masculini^ and nationhood
in heavy metal scenes in Norway, South Africa and Australia
Catherine Hoad
Abstract of a thesis for a Doctorate of Philosophy submitted to Macquarie University, Sydney, Australia
In response to the current dynamics of the
Global Metal model, this thesis draws atten¬
tion to how whiteness maintains an invisibi-
lised instrumental hegemony within heavy
metal music, even as research on the genre
continues to grow. I address the complex
problem of how whiteness is represented in
heavy metal scenes and practices, both as a
site of academic inquiry and force of cul¬
tural significance. I argue that the whiteness
and white heteromasculinity of heavy metal
emerges in disparate locales as expressions of
distinct nationalist projects. This research
addresses the national specificity with which
whiteness is valorised in heavy metal scenes,
and how disparate national identities are
tacitly and explicitly tied to white hetero¬
masculine identity.
This thesis negotiates scholarly ways of
addressing whiteness in heavy metal that
move beyond discussions of demographics,
virtuosity and spectacular racism. I analyse
how the normalisation, construction and
performance of whiteness, masculinity and
nationhood within heavy metal scenes can
have profound, pervasive and systematic
oppressive consequences. The objective
of my thesis is therefore to unveil the (in)
visibility of whiteness within heavy metal
scenes, and indicate how such whitenesses
are deployed within particular countries, as
both explicit political violence and instru¬
mental hegemony. The quest in pointing to
the fragmentation and multiplicity of white¬
nesses across three different countries is to
deconstruct the structure of white hegemony,
and call into question the strategic political
position that emerges in treating white selves
as a uniform category.
This research maps the matrix of white¬
ness, masculinity and nationhood through
which heavy metal scenes across Norway,
South Africa and Australia have produced
and defended national identity. I distinguish
three key forms of white nationalism- — Nor¬
way’s monstrous nationalism, in which the
nation is constructed as terrifying and atavis¬
tic; South Africa’s resistant nationalism, which
responds to post-Apartheid claims of white
victimhood; and Australia’s banal nationalism,
which consecrates mundanity as an authentic
national condition. Such constellations of
whiteness, masculinity and nationhood have
enabled tacit and explicit constructions of
exclusionary communities formed through
collective memory and territory. These scenes
are demonstrative of the ways in which white
inflections inform the practices of both heavy
metal scenes and the specifically local white¬
nesses manifest within them.
Dr Catherine Hoad,
Department of Media, Music, Communica¬
tions and Cultural Studies,
Macquarie University,
Sydney NSW 2109
AUSTRALIA
Email: catherine.hoad@mq.edu.au
94
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
p. 95. ISSN 0035-9173/16/010095-01
Thesis abstract
Interhemispheric asymmetry of global warming:
the role of ocean dynamics
David Karel Hutchinson
Abstract of a thesis for a Doctorate of Philosophy submitted to the University of New South Wales,
Sydney, Australia
Global surface air temperature is increasing
due to rising greenhouse gases. This warming
has occurred at a faster rate in the Northern
Hemisphere (NH) than the Southern Hemi¬
sphere (SH) and the asymmetry of warming
between hemispheres is predicted to con¬
tinue throughout the 2 1 st Century. Several
factors contribute to this interhemispheric
asymmetry, including the greater propor¬
tion of land in the NH and the northward
transport of heat by the ocean. This thesis
focuses on the role of ocean dynamics in set¬
ting the warming asymmetry, using several
modelling approaches.
First, the impact of the Antarctic Cir¬
cumpolar Current (ACC) on the interhemi¬
spheric warming asymmetry is investigated.
The role of the ACC is isolated by comparing
warming experiments in a global coupled
climate model with and without a land bar¬
rier across Drake Passage (DP). With DP
closed, the asymmetry in sea surface tem¬
perature (SST) warming is reduced, due to
the presence of a subpolar gyre, and a lower
Antarctic sea ice extent.
Second, the asymmetry of warming is
examined when moving from coarse (1°) to
eddy-permitting (0.25°) ocean resolution.
We use an idealised coupled model with a
60° sector ocean domain, comprising one
basin with Atlantic-like bathymetry and an
ACC channel. A larger high latitude SST
asymmetry develops in the 0.25° model than
the 1° model, both in control runs and in
warming scenarios. The larger warming
asymmetry in the 0.25° model is caused by
stronger boundary current heat transport
and reduced NH sea ice. The SH warming is
less sensitive to the resolution change, since
eddy heat transport differences between
the models are small compared with mean
flow heat transport differences. When SH
westerly winds are enhanced, the warming
asymmetry increases, with greater upwelling
of cool water in the Southern Ocean and
greater warming in the NH.
Finally the impact of realistic bathymetry
is explored in the sector climate model. The
Atlantic-like sector model is compared with
a flat bottom rectangular model in similar
experiments. The Atlantic and rectangular
models have similar control climates, how¬
ever the rectangular models have a stronger
subpolar gyre in the NH in the absence of
bathymetry. In warming experiments, the
rectangular models develop warming and
cooling regions in the NH, while the Atlantic
models have no significant cooling regions.
The Atlantic models exhibit greater sensitiv¬
ity of ACC transport to wind forcing.
Dr David Hutchinson,
Climate Change Research Centre,
University of New South Wales,
Sydney NSW 2052 AUSTRALIA
Email: david.hutchinson@geo.su.se
Thesis: handle.unsw.edu.au/ 1 959.4/55380
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p. 96. ISSN 0035-9173/16/010096-01
Thesis abstract
Conscious and not-conscious processing of
visual mismatch negativity
Bradley N. Jack
Abstract of a thesis for a Doctorate of Philosophy submitted to Southern Cross University, Lismore,
Australia
The general aim of my thesis is to investi¬
gate conscious and not-conscious process¬
ing of sequences of stimuli that yield
visual mismatch negativity (vMMN), a
well-established brain signature of predic¬
tion and prediction-error. vMMN is typi¬
cally observed in the oddball paradigm: an
infrequent visual stimulus— a deviant, is
randomly and unpredictably presented in a
sequence of more frequent visual stimuli —
the standards. vMMN is a negative compo¬
nent of event-related potentials (ERPs), and
is seen most clearly in the difference wave:
the ERP for the deviant minus the ERP for
the standard, between 150 and 400 ms after
stimulus onset.
To investigate conscious and not-con¬
scious processing of vMMN, I conducted
four electroencephalography (EEG)/ERP
experiments. In Experiment 1, I showed
that it is easier to find neural correlates of
visual consciousness — differences in brain
activity between conscious and not-con¬
scious visual stimuli, with cardinal gratings
than with oblique gratings. In Experiment 2,
I showed that a source of information about
which we are not-conscious, eye-of-origin
(utrocular) information, yields a reliable
vMMN. In Experiment 3, 1 hid my deviants
from visual consciousness using binocular
rivalry suppression, and found that the size
of vMMN is smaller to that elicited by the
same stimulus when it is conscious during
binocular rivalry dominance. In Experiment
4, I hid my standards and deviants from
visual consciousness using continuous flash
suppression (CFS), and found that the size
of vMMN is bigger than that elicited by the
same stimuli when they are conscious.
My results are consistent with the notions
that our brains establish predictive models
of visual perception about regular visual
input, that our brains are constantly testing
the reliability of these models, and that our
brains update these models when something
unexpected occurs. My results also show that
these processes are independent of visual
consciousness. I conclude that visual con¬
sciousness is not necessary to elicit vMMN,
confirming that vMMN is an automatic
brain response.
Dr Bradley N. Jack
School of Health and Human Science
Southern Cross University
Lismore NSW 2480
AUSTRALIA
Email: bradley.jack@unsw.edu.au
96
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p. 97. ISSN 0035-9173/16/010097-01
Thesis abstract
White matter microstructural decline and
cognitive performance in older adults:
the influence of cardiovascular health
Todd Jolly
Abstract of a thesis for a Doctorate of Philosophy submitted to University of Newcastle, Newcastle,
Australia
Age-related cognitive decline is well docu¬
mented, especially in memory, speed of
processing and executive functions. Struc¬
tural brain changes are also well documented
but often do not directly map onto the mild
cognitive decline seen in otherwise healthy
older adults. Recent work has focused on
whether cognitive ageing is associated with
decline in the strength of structural con¬
nectivity between neural regions, using
diflhision magnetic resonance imaging
(dMRI) . Reduced integrity of white matter
microstructure across the whole brain and
in regions of interest, as measured by frac¬
tional anisotropy (FA), has been shown to
be associated with cognitive decline in older
adults who show no signs of dementia. This
thesis uses dMRI tractography to examine
the association between multiple measures of
white matter microstructure across the whole
brain and in 18 major white matter tracts
and cognitive performance on a range of
tasks that vary in process specificity. Seventy
non-demented older adults (aged 43-87y)
with varying degree of white matter disease
completed a comprehensive cognitive and
imaging assessment. Cognitive functioning
was assessed at three levels: Firstly, global
cognitive functioning was assessed using the
Montreal Cognitive Assessment (MoCA).
Then through the use of standardised neu¬
ropsychological tests, more specific cogni¬
tive domains of working memory, episodic
memory, executive function and processing
speed were assessed. An experimental task
switching paradigm was then used to assess
more specific components of executive func¬
tion relating to proactive and reactive con¬
trol processes. These showed that ability to
detect the impact of tract-specific changes
in white matter microstructure on cognitive
performance was dependent on the specifi¬
city of the cognitive test. Although, irrespec¬
tive of the level of cognitive assessment, the
relationship between decline in white matter
microstructural integrity and cognitive per¬
formance was specific only to participants
with poor cardiovascular health. These find¬
ings suggest that cognitive and brain ageing
profiles in older adults vary as a function of
cardiovascular health and have strong impli¬
cations for theories of cognitive ageing. They
also emphasise the importance of cardiovas¬
cular health in prevention or delay in onset
of cognitive decline in old age.
Dr Todd Jolly,
School of Psychology,
University of Newcastle,
Newcastle NSW 2300
AUSTRALIA
Email: Todd.Jolly@uon.edu.au
97
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pp. 98-99. ISSN 0035-9173/16/010098-02
Thesis abstract
Achieving change in student assessment in Vietnamese
teacher training institutions
Gam Thi Hong Luong
Abstract of a thesis for a Doctorate of Philosophy submitted to Southern Cross University, Lismore,
Australia
The role of student assessment in shaping
learning outcomes is well established. In
Vietnam, there is a developing consensus at
official levels that reform of student assess¬
ment practices in higher education institu¬
tions is required. In 2006 and 2007, the
Ministry of Education and Training issued
Decisions seeking to encourage higher edu¬
cation institutions to make more use of stu¬
dent assessment methods likely to support
activity-based and self-directed approaches
to learning. To date, however, Vietnamese
universities and colleges have been remark¬
ably slow to respond. They continue to rely
on traditional standardised tests that pro¬
mote rote learning and do little to develop
critical thinking or problem-solving skills
among learners.
The present investigation seeks to provide
an understanding of the beliefs, values and
attitudes towards student assessment of a
group of lecturers and educational managers
from three teacher training universities in
Vietnam. Its purpose is to throw light on the
conditions affecting their ability and willing¬
ness to reform student assessment practices
at their institutions. Theoretical perspec¬
tives on student assessment from empiri¬
cal research in developed higher education
systems inform the investigation, and three
theories of educational change are drawn
upon in seeking to identify the factors that
might impact on the student assessment
reform process in higher education institu¬
tions in Vietnam.
An ethnographic approach is taken to the
collection of data, and Naturalistic Inquiry
(Lincoln & Cuba, 1985) provides a meth¬
odological framework for the investigation.
Ethnographic interviews were conducted
with 24 experienced members of academic
staff from across the three site institutions.
TEese participants were selected using a
‘snowbair sampling technique whereby each
was recommended by a colleague as being
interested in and experienced with issues in
student assessment. The interview data were
analysed by means of thematic analysis, with
particular regard taken to ensure the trust¬
worthiness of the findings.
Three distinct groups of participants are
identified. For three of the participants, atti¬
tudes to teaching and student assessment
were strongly teacher-centred, supportive of
traditional standardised methods of student
assessment, and shaped by beliefs that stu¬
dents should be obedient, passive learners.
These participants had a limited understand¬
ing of the range of approaches to student
assessment: they were unwilling to make
any changes in terms of how they assessed
student learning. For 13 of the participants,
however, there was recognition of the need
to reform student assessment practices:
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Journal & Proceedings of the Royal Society of New South Wales
Luong— Change in Student Assessment
these participants expressed a willingness
to change their own assessment practices,
but they felt constrained from doing so
because of a perceived lack of expertise and
because they saw that many more hours of
work would be required to do so effectively
This group, therefore, had not implemented
any significant changes. The third group
of eight participants aspired to reform the
ways in which students were assessed: they
actively implemented measures intended to
achieve effective reform. They were more
inclined than any of the other participants
to value their students as learners. They also
claimed to be strongly supportive of the role
and importance of formative assessment.
The investigation points to the importance
of achieving an alignment between policy,
leadership and practice in order to achieve
enduring educational change. This align¬
ment requires persistent effort to be directed
at ensuring that all relevant stakeholders
are properly informed about the goals and
objectives of desired change. It also requires
them to have the resources needed to engage
meaningfully in the change process by imple¬
menting continuous assessment and forma¬
tive feedback to learners about their learning
progress. They must also have opportunities
to converse collaboratively with their peers
about why and how assessment practice
needs to be improved.
Achieving a more enlightened approach
to student assessment on a national scale in
Vietnam’s higher education system appears
for the time being to remain a distant pros¬
pect. This investigation does, however, pro¬
vide insights into what might need to be
done to make the aspiration more achievable,
more rapidly.
Dr Gam Thi Hong Luong
School of Education
Southern Cross University
Lismore NSW 2480
AUSTRALIA
Email: gamluong2012@yahoo.com
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p. 100. ISSN 0035-9173/16/010100-01
Thesis abstract
Zone of impeachment: a post-Foucauldian analysis of
controlled operations law and pohcy
Brendon Murphy
Abstract of a thesis for a Doctorate of Philosophy submitted to University of Newcastle, Newcastle,
Australia
This thesis presents a Foucauldian analy¬
sis of Australian controlled operations law.
The purpose was to extend current doctrinal
scholarship by exploring the discursive forces
that shape this highly invasive and contro¬
versial investigative power. This thesis con¬
tends that the present doctrinal understand¬
ing is incomplete, and largely unaware of the
epistemological forces operating within law
and policy. By deploying a Foucauldian ana¬
lytic we can extend our understanding of the
complex relationship between knowledge
systems, discourse, power and law.
Through the deployment of a nomadic,
grounded genealogy in the analysis of con¬
trolled operations Second Reading Speeches,
this research found that the governing ration¬
alities of controlled operations law and policy
is linked to an imperative logic dominated
by discourses of risk, audit and exceptions.
This dynamic explains why controlled opera¬
tions legal architecture and policy is in its
current form. Far from being a reaction to
the decision in Ridgeway, controlled opera¬
tions law is part of a legal and cultural shift
in law enforcement, characterised by com¬
plex relationships between risk, rights, law
and citizenship. The controlled operation is
revealed as a form of apparatus: a technology
of truth and power, facilitated by law.
This insight allows us to re-imagine the
relationship between law, rights, citizenship
and sovereignty in late modernity. In this
environment the investigative apparatus of
the controlled operation creates a field of
governance within the private space of liberal
citizenship, revealing the true character of
citizenship in late modernity as a zone of
impeachment ■” a location in which rights
are fragile and open to perpetual potential
derogation and modification. In this zone
the rights attached to liberal conceptions
of citizenship are increasingly the subject
of subordination to a risk imperative and a
logic of exception.
Dr Brendon Murphy,
Department of Business and Law,
University of Newcastle,
Newcastle NSW 2300
Australia
Email: brendon.murphy@newcastle.edu.au
100
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 101-102. ISSN 0035-9173/16/010101-02
Thesis abstract
The effects and mechanisms of the therapeutic hypothermia
on intracranial pressure regulation following ischaemic
stroke in rats
Lucy Mujrtha
Abstract of a thesis for a Doctorate of Philosophy submitted to University of Newcastle, Newcastle,
Australia
Background: Intracranial pressure (ICP) rises
to dangerous levels 2 to 5 days after large
ischaemic stroke. ICP following small stroke
is not routinely monitored, although animal
data suggests ICP rises 24 hours following
small experimental stroke. Cerebral oedema
has been thought to be the primary cause for
ICP elevation. This assumption may have
risen because ICP has only been monitored
in patients with large infarct and oedema
volumes. Since small ischaemic infarcts
cause less cerebral swelling, ICP elevation
may be the result of a different mechanism(s).
Recent human imaging data indicates that
patients deteriorating soon after minor
stroke do so on the basis of cerebral col¬
lateral blood flow failure. Until now there
has not been a plausible explanation for this
'collateral failure”. Long-duration hypother¬
mia has been shown to lower ICP in patients.
Long durations of cooling increase the risk of
infection and rebound ICP during rewarm¬
ing. Short-duration hypothermia has shown
overwhelming efficacy in animal models of
stroke but has not been tested in humans. I
hypothesise: that ICP increases at 24 hours
after small stroke; that this rise is not due to
cerebral oedema; that ICP elevation reduces
collateral blood flow; and that short-duration
moderate or mild hypothermia prevents ICP
elevation post-stroke. Methods: An epidural
ICP monitoring technique was developed.
Experimental ischaemic stroke (middle cer¬
ebral artery occlusion) was performed in
Long Evans, outbred Wistar and Sprague-
Dawley rats and ICP was monitored. Infarct
and oedema volumes were calculated using
wet-dry weight calculations, histology or in
vivo magnetic resonance imaging. Collat¬
eral blood flow was visualized using fluores¬
cent microspheres through a closed cranial
window and recorded using a high-speed
microscope-mounted recording camera.
Short-duration moderate (32.5°C) or mild
(35°C) hypothermia, or normothermia
(37°C) was administered 1 hour post-stroke.
Results: Mean ICP was 9.1 ± 5.2 mm Hg at
baseline (pooled - all animals). ICP was sig¬
nificantly elevated 24 hours post-stroke in all
normothermic animals (40.3 ±16 mm Hg,
pooled normothermic animals,/? < 0.0001
vs. baseline). Mean infarct volume was
22.6 ± 17.5% of contralateral hemisphere.
Oedema volumes were small and were not
correlated with ICP post-stroke (r^ = 0.09,
p = 0.15). There was a strong correlation
between ICP elevation and collateral blood
flow decrease (r = -0.62,/? < 0.0001). Early
intervention of short-duration hypothermia
completely prevented ICP rise post-stroke
(10.3 ±6.5 mm Hg, pooled hypothermic
animals at 24 hours,/? < 0.0001 vs, normo-
101
Journal & Proceedings of the Royal Society of New South 'V^es
Murtha — Effects of TJierapeutic Hypothermia
thermic animals at 24 hours). Conclusions:
In this thesis, I have presented data that con¬
tradicts the accepted wisdom in several ways
and has important implications for patients
with stroke. It suggests that ICP could be
elevated in patients with small stroke and
that a factor other than oedema is the pri¬
mary cause of this ICP elevation. The data
also suggest that ICP elevation following
stroke is the likely mechanism of collateral
failure leading to neurological deterioration
in stroke patients. Finally, I have demon¬
strated that short-duration hypothermia is
an effective ICP preventative treatment fol¬
lowing experimental stroke, and suggests that
short-duration hypothermia clinical stud¬
ies in humans is warranted. These findings
suggest that a fundamental rethink of ICP
regulation post-stroke is necessary and have
potentially important and exciting implica¬
tions for the future treatment of stroke and
stroke-in-progression.
Dr Lucy Murtha,
School of Biomedical Sciences and Pharmacy,
University of Newcastle,
Newcastle NSW 2300
AUSTRALIA
Email: Lucy.Murtha@newcastle.edu.au
102
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
p. 103. ISSN 0035-9173/16/010103-01
Thesis abstract
Palaeontolog)^, taxonomy and biostratigraphy of Cambrian
assemblages from the Pertaoorrta Group, Amadeus Basin,
Northern Territory
Patrick Mark Smith
Abstract of a thesis for a Doctorate of Philosophy submitted to Macquarie University, Sydney, Australia
The Amadeus Basin is a large sedimentary
province in central Australia that covers an
approximate area of 170,000 square kilome¬
tres. Despite the known occurrence of fos¬
sils from the majority of stratigraphic units
within the Cambrian Pertaoorrta Group
there is a dearth of published palaeonto¬
logical data, including no comprehensive
biostratigraphy.
Presented as part of this thesis is a detailed
investigation into three formations span¬
ning the Cambrian Series 2“3 units of the
Pertaoorrta Group. The oldest of these, the
Tempe Formation and Giles Creek Dolos-
tone have previously been regarded as coeval.
Examination of specimens from both drill-
core and outcrop material from these two
formations revealed a considerable diversity
of new and biostratigraphically-informative
fossils. The described taxa provide evidence
that these two sedimentary units were depos¬
ited at different times. The Tempe Formation
(in Paper 1) belongs to the Ordian, whereas
the fauna from the Giles Creek Dolostone
(in Papers 2—4) is distinctly younger and cor¬
relates with the overlying early Templetonian.
These results suggest that the current regional
stratigraphic scheme needs to be amended.
The youngest stratigraphic unit examined
in this thesis is the Goyder Formation (in
Paper 5). The initial age estimates for this
formation were based solely on vague reports
of trilobites. Our collections demonstrate
that the Goyder Formation contains a highly
diverse fossil fauna with at least 20 different
trilobite taxa. This assemblage indicates a
late Mindyallan age (equivalent to Cambrian
Series 3, Guzhangian) within the Glyptag-
nostus solidotus Zone.
Detailed logging and sampling through
formations in the Pertaoorrta Group has
allowed for precise ages where little to no
biostratigraphic data had previously been
available. These ages have facilitated the
development of a preliminary quantitative
biostratigraphy of the Gambrian Series 2-3
portion of the Amadeus Basin, thus permit¬
ting more accurate intra- and interbasinal
correlation.
Patrick Mark Smith,
Department of Biology,
Macquarie University,
Sydney NSW 2109
AUSTRALIA
Email: Patrick.mark.smith. 1 990@gmail.com
103
Journal & Proceedings of the Royal Society of New South Wales, vol. 149, parts 1 & 2, 2016,
pp. 104-105. ISSN 0035-9173/16/010104-02
Thesis abstract
Targeted, one-to-one instruction in whole-number
arithmetic: a framework of key elements
Thi Le Trln
Abstract of a thesis for a Doctorate of Philosophy submitted to Southern Cross University, Lismore,
Australia
In Australia, although there has been strong
advocacy for individualised intervention
programs, there is a limited research litera¬
ture available that focuses on teacher-student
interactions and teaching practices related to
one-to-one instruction. This investigation
seeks to address that gap. Its aim is to iden¬
tify and illuminate the nature of Key Ele¬
ments of one-to-one instruction that expert
tutors use when interacting in intensive,
one-to-one instruction of whole-number
arithmetic with Years 3 and 4 students. A
Key Element is a micro-instructional strategy
that is the smallest unit of analysis of highly
interactive one-to-one instruction.
The investigation draws on data collected
within the framework of the Mathematics
Intervention Specialist Program (Wright,
Ellemor-Collins & Lewis, 2011). From
this source, approximately 33 hours of video
recordings of teaching sessions involving four
teachers and six students were analysed.
The theoretical perspective underpinning
the investigation is interpretative. Within
this perspective, a phenomenological
approach was used to gain insight into the
essence of the Key Elements of one-to-one
intervention teaching. A standard method
for analysing the data, that is, ‘close observa¬
tion” (Van Manen, 1997, p. 68), in which
the Key Elements are viewed as the central
phenomenon requiring exploration and
understanding, was employed. The analytical
techniques described by Van Manen (1990,
1997), and further elaborated as procedures
for phenomenological analysis by Hycner
(1999), were applied. As well, the investi¬
gation utilised methodological approaches
described by Cobb and Whitenack (1996),
and by Powell, Francisco, and Maher (2003),
for analysing large sets of video recordings.
Twenty-five Key Elements were identified
and, for each, a deeply layered description
was developed. As well, a comprehensive
framework for analysing one-to-one instruc¬
tion was conceptualised. The framework
shows how Key Elements can be used to
analyse intensive, one-to-one instruction in
whole-number arithmetic.
The investigation advances understand¬
ing about teacher-student interactions and
teaching practice in intensive, one-to-one
interventions. Understanding the Key
Elements leads to more effective ways to
characterise the instructional strategies that
teachers utilise in one-to-one intervention
teaching. The framework developed con¬
stitutes an extension of the current body of
theoretical knowledge about targeted one-to-
one intensive intervention in whole-number
arithmetic. It will inform teachers who are
working with low-attaining students by
104
Journal & Proceedings of the Royal Society of New South Wales
Tran— -Instruction in Whole-Number Arithmetic
providing useful information about teacher-
student interaction in mathematical inter¬
ventions, which in turn may illuminate how
particular teaching intervention practices
influence student learning outcomes (Tran
& Wright, 2014b).
Dr Thi Le Tran
School of Education
Southern Cross University
Lismore NSW 2480
AUSTRALIA
Email: tranlethicdsp(2)yahoo.com
105
Proceedings of the Royal Society of New South Wales
The 2016 programme of events — Sydney
Held at the Union, Universities and Schools Club, 25 Bent St, Sydney unless otherwise stated.
Wed 3 Feb
1240^^ Ordinary
Meeting
RSNSW
Scholarship
winners
Adrian Dudek
Charles Forster
Yevgeny Stadnik
Australian National
University
University of Sydney
University of New South
Wales
Mon 25 Feb
The Four
Societies Lecture
Professor Robert Clark AO
FAA Dist FRSN
Chak of Energy Strategy and
Policy, University of New
South Wales
Australian Energy Policy
Held in conjunction with the Nuclear Engineering Panel of the Sydney Branch of
Engineers Australia, the Australian Nuclear Association and the Australian
Institute of Energy.
Held at Hamilton and Parkes Rooms, Level 47, MLC Centre, King and
Castlereagh St.
Wed 4 Mar
124Lt Ordinary
Meeting
Dr Len Fisher
Visiting Fellow in Physics,
University of Bristol
How to win an IgNobel Prize
and other adventures in
communicating science
Wed 16 Mar
Joint Lecture of
the Australian
Institute of
Physics and the
RSNSW
Professor Ron Grunstein
Woolcock Institute of
Medical Research,
University of Sydney and
Royal Prince Alfred Hospital,
Head
From Snoring to
Somnambulism —
The Mystery of the Sleeping
Brain
Joint Meeting with
School, Latham T
L the Australian Institute of Physics, held at Trinity Grammar
leatre, 119 Prospect Road, Summer HiU
Wed 6 April
1242^^ Ordinary
Meeting and
149* Annual
General Meeting
Dr Donald Hector FRSN
President of the Royal Society
of New South Wales
Presidential address: Royal
Society of NSW — relevance
in the 21st century
'i;
Wed 4 May
Annual Dinner:
Distinguished
Fellow's Lecture
and presentation
of the Society's
2016 awards
Guests of honour: The
Society's Vice-Regal Patron,
Flis Excellency General The
Honourable David Hurley AC
DSC (Ret'd), Governor of
New South Wales and
Em. Professor Eugenie
Lumbers AM DistFRSN
Science Policy and
University Research
j Wed 1 June
1243^^ Ordinary
Meeting
Professor Peter Hiscock
Tom Austen Brown Professor
of Australian Archaeology,
University of Sydney
The curious case of the
scientist in cinema: how
Indiana Jones turns out to
be the bad guy!
Wed 6 July
1244th Ordinary
Meeting
Dr Bob Young
Associate Professor of
Geoscience (ret’d), University
of Wollongong
"Royal" not "Philosophical"
- W.B. Clarke's Inaugural
Address to the Royal Society
of NSW
Wed 3 Aug
1245* Ordinary
Meeting
Dr. Barbara Briggs, Honorary
Research Associate Royal
Botanic Gardens
Celebrating the 200*
Birthday of Royal Botanic
Gardens: A Personal
History of 57 years of
Science
Sydney Science Festival lunchtime science talks
Fri 12 Aug
Sydney Science
Festival
Professor Mikhail
Prokopenko, University of
Sydney
Complex Systems and
Swarm Intelligence
Tu 1 6 Aug
Sydney Science
Festival
Dr Brett SummereU
Royal Botanic Gardens
The Royal Botanic Gardens
200th Birthday
,Wedl7Aug
Sydney Science
Festival
Em Professor Br^mn Plibbert
UNSW, President RSNSW
Courts, Criminals and
Chemistry: Forensic Science
in NSW
Thu 1 8 Aug
Sydney Science
Festival
Professor Pascal Perez
University of Wollongong
Community-driven Internet
of Things: the new
revolution?
Wed 7 Sep
1246* Ordinary
Meeting
Richard NeviQe
Ivlitchell Dbrarian and
Dkector, Education &
Scholarship State Dbrary of
NSW
A source of inspiration and
delight: The Mitchell Library
Wed 5 Oct
1247* Ordinary
Meeting
Professor Itai lanev
School of Engineering,
University of Sydney
From sand and rice bubbles
to earthquakes and
volcanoes
Thu 13 Oct
2016 Dirac
Lecture
Duffield Professor Kenneth
Freeman
Australian National University
Dark Matter in the Universe
Wed 2 Nov
1248* Ordinary
Meeting
Professor E. James Kehoe
Professor of Psychology,
UNSW
Finding the Right Course
for the Right Horse: Recent
Evidence-Based Advances
in Instructional Design
Tue 29 Nov
RSNSW and
Four Academies
Forum
Government House, Sydney;
hosted by his Excellency
General The Honourable
David Hurley AC DSC (Ret’d)
Governor of NSW and Patron
of the Royal Society of NSW
at Government House
Society as a complex system:
Implications for Science,
Practice and Policy and
Celebration of the 1 50*
Anniversary of Royal
Patronage
Held in cooperation with the Australian Academy of Science, the Australian
Academy of Technological Sciences and Engineering, the Australian Academy of
the Humanities and the Academy of Social Sciences in Australia.
Wed 7 Dec
1249* Ordinary
Meeting
Royal Society of NSW 2016
Jak Kelly Award:
Matthew Barr
School of Mathematical and
Physical Science, Newcastle
University
Imaging with a deft touch —
The scanning helium
microscope
The 2015 programme of events -- Southern Highlands
Held at the Performing Arts Centre, Chevalier College, Bowral.
Thu 18 Feb
Dr Charley Lineweaver
School of Astronomy & Astrophysics
and Earth Sciences, Australian
National University
Death and Nothingness: Why did
Death Evolve? Why is there Something
Rather than Nothing?
Sat 12 Mar
Dr Christian Heim and Dr CaroHne
Heim
Special Event: An Afternoon with
Chopin and George Sand
Thu 21 Apr
Prof Gordon Parker
Scientia Professor of Psychiatry,
UNSW, Executive Director of the
Black Dog Institute
Winston Churchill, bipolar disorder, and
the Dardanelles campaign
Thu 1 9 May
Dr Kathleen Riley
Freelance writer, theatre historian and
critic
The Science of Spontaneity: Fred
Astaire as the Consummate Craftsmen
Thu 16 June
Dr Ken McCracken
Solar physicist, foundation director of
CSIRO Office of Space Science and of
CSIRO Division of Mineral Physics
The Sun, Sunspots, and Space Weather
Thu 21 July
Dr David Suhy
Chief Scientific Officer, Benitec
Biopharma
Silencing Genes for Life
Thu 1 8 Aug
Ian Skinner
Neuroscience Research Australia
(NeuRa)
Chronic Pain
Thu 1 5 Sep
Assoc Prof Tony Masters
Chair of the Academic Board,
University of Sydney
Sustainability - Chemical Solutions for a
Tricky Problem
Thu 20 Oct
Prof Gordian Fulde
Director of emergency at St Vincent’s
Hospital, Senior Australian of the Year
2016
Do we have a problem? — Hospital
emergency, alcohol and drugs
Thu 17 Nov
Prof Gordon Wallace
Director, ARC Centre of Excellence
for Electromaterials Science,
University of WoUongong
BioPrinting: 3D Printing Parts for
Bodies
f Awards for 2016
The Council of the Royal Society of New South Wales have determined to make the following awards for
2016:
1. Bdgeworth David Medal.
Associate Professor Simon Ho
ARC Queen Elizabeth II FeUow, School of Biological Sciences, University of Sydney.
I The Edgeworth David Medal, established in memory of Professor Sir Tannatt William Edgeworth David FRS,
a past President of the Society, is awarded for distinguished contributions by a young scientist under the age of
35 years.
Associate Professor Simon Ho has made hugely important contributions and developed new methods in the
field of ‘molecular clocks' in biology ~ a way of estimating evolutionary rates and timescales from DNA
sequences using statistical models. These estimates underpin a broad range of studies in conservation genetics,
speciation and diversification, domestication of animals and plants, events in human prehistory, and the
population dynamics of pathogens. His research focuses on understanding how evolutionary rates vary at the
: genomic level and estimating the timescale of the Tree of Life. These are fiindamental goals of biological
: inquiry because they deal with the details of the evolutionary process.
I Simon Ho’s work has set a range of standards in the field, as well as producing methods and practises that are
now widely used by researchers. His research has led to important improvements in the way that researchers
: estimate evolutionary timescales using genetic and genomic data, with significant consequences for our
understanding of the evolutionary past.
2. History and Philosophy of Science Medal
i.
5 Professor Warwick Anderson
! ARC Laureate Fellow and Professor in the Department of History and the Centre for Values, Ethics and the
I Law in Medicine, University of Sydney. Additionally, he has an affiOiation with the Unit for History and
I Philosophy of Science at Sydney University.
[ The Royal Society of NSW History and Philosophy of Science Medal was established in 2015 to recognise
t outstanding achievement in the Idistory and Philosophy of Science. The medaUist will have made a significant
i contribution to the understanding of the history and philosophy of science, with preference being given to the
I study of ideas, institutions and individuals of significance to the practice of the natural sciences in Australia.
Professor Anderson is a medical doctor turned historian, who has made important contributions to the history
of science, medicine, and pubHc health; the history of racial thought and postcolonial science studies. He is a
Fellow of the Academy of the Social Sciences, and won the 2014 History of Science Society's Price/Webster
Prize for the article, “Hybridity, Race and Science: The Voyage of the Zaca, 1934-1935."
In 2014 Professor Anderson, with immunologist Ian Mackay, wrote a brilliant and original book. Intolerant
Bodies: A Short Histoty of Autoimmunity^ published by Johns Hopkins University Press. The authors foUow the
puzzle of autoimmunity from theory to laboratory practice to individual patients' case histories. The result is a
compelling study of concepts in action. This sophisticated but highly readable history helps close the gap
between medical science and the general public's understanding.
3. Clarke Medal for Zoolog)j.
Professor Christopher Dickman
University of Sydney School of Biological Sciences.
The Clarke Medal was established to acknowledge the contribution by Rev William Branwhite Clarke MA FRS
FGS, Vice-President of the Royal Society of New South Wales from 1866 to 1878. The Medal is awarded
annuaQy for distinguished work in the natural sciences of geology, botany and zoology done in Australia and its
Territories.
Professor Dickman’s major contributions He in terrestrial zoology and ecology. He has long been curious
about the factors that promote and maintain biodiversity, especially among land mammals and other terrestrial
vertebrates. For the last 35 years he has focused in particular on understanding the forces that shape the
distribution and abundance of AustraHa’s endemic mammals and identifying the factors that are causing so
many species to decHne. His ground-breaking work on AustraHa’s desert mammals and on the continent's
Hitroduced predators have gained him a formidable reputation as a leading national and international authority
on mammaHan ecology.
4. Royal Society of New South Wales Scholarships.
Adrian Dudek (AustraHan National University, School of Mathematics)
Yevgeny Stadnik (University of New South Wales, School of Physics)
Charles Foster (University of Sydney, School of Botany)
The Council of the Society firnds the Royal Society of New South Wales Scholarship in order to acknowledge
outstanding achievements by early-career individuals working, in a science-related field within New South
Wales or the AustraHan Capital Territory, towards a research degree in a science related field.
Adrian Dudek is working in number theory under Dr Trudgian at the ANU. During his PhD he has
pubHshed (or had accepted) eight papers in the peer reviewed Hterature. Plis appHcation explained his research
thus: “In particular, Tm interested in the elusive tale of the prime numbers. When I let this sHp to most
people, somewhat tepid memories of their primary school days are horrificaUy conjured. However, the prime
numbers have been studied for thousands of years, or at least since 300BC, when the great Greek geometer
EucHd proved that tliere are infinitely many of them. Since such ancient times, the primes have attracted the
attention of curious mathematicians (and other characters) for one reason: it’s extraordinarily difficult to
understand the behaviour of the prime numbers. For Histance, if you were to write down a Hst of the first 100
prime numbers (a rousing exercise for a Friday night, Tm sure!), you would not be able to find an intelHgible
pattern. That being said, some recent spectacular advances in number theory mean that the prime numbers are
becoming less elusive and more understandable ...”
Yevgeny Stadnik works with Professor Flambaum FRSN on “Manifestations of Dark Matter and Variation
of Fundamental Constants in Atoms and Astrophysical Phenomena”. He writes: “My project is on the
investigation of new effects produced by dark matter and proposing novel ways of detecting dark matter. We
have pubHshed a number of important works in this direction, including results that already improve on
existing sensitivities in the detection of certain types of dark matter by up to 15 orders of magnitude. Our
results have been pubHshed in leading physics journals, including three pubHcations in Physical Review Letters
(which is the most highly cited physics journal), and have contributed to the initiation of a number of new
laboratory searches worldwide.”
Charles Forster is a botanist working with our Edgeworth David medaUist Simon Ho on a project “Using
genome-scale data to untangle the evolutionary history of flowering plants”. A University of Sydney medalHst,
Charles has been able to estimate the timescale of evolution of a range of plants using genomic data. His
analyses have been careful and comprehensive, and he is on the verge of pubHshing his outstanding work on
this topic. This is in addition to three papers from his honours research and three pubHshed or under review.
This work has also led to the development of some important research coUaborations with coUeagues at the j
Royal Botanic Gardens (Sydney) and Universite Paris-Sud (France). He writes: “I have provided the most
comprehensive combination of analyses of the angiosperm evolutionary timescale so far. The results I have
obtained reflect the increasingly common finding that molecular dating estimates predate the oldest fossils by a
non-trivial amount of time, up to 70 mOHon years when considering mean estimates.”
5. The Royal Society of New South Wales and A.ustralian Institute of Physics Jak Kelly A.ward.
James CoUess
University of Sydney, School of Physics
The Jak Kelly Award is awarded joindy with the Australian Institute of Physics (AIP) to the best PhD
student talk, this year presented to a joint meeting with the AIP held on November 17 at Trinity
Grammar School.
James CoUess is a postgraduate student at the University of Sydney currently undertaking his PhD under the
supervision of Professor David ReiUy. His research focus is readout and control techniques for GaAs spin
qubits. James hopes his research wiU influence the design and fabrication of reliable multiqubit gates. His talk
was entitled “From Quantum Devices to Quanmm Machines”. It explored the complexity of scaling quantum
processors and discussed new techniques and hardware developed to meet these chaUenges. In particular,
James had developed new methods of readout that aUow the dispersive sensing of single-electrons using
integrated sensors and the capability to read out multiple qubits simultaneously. A scalable control scheme is
also demonstrated allowing large numbers of qubits to be manipulated with a small number of input signals.
The award consists of an engraved plaque, a $500 prize and a year's membership of the Society. As the winner
of the Jak KeUy award, James presented his talk to Royal Society on the of November at the Union,
Universities and Schools Club.
Archibald Liversidge:
Imperial Science under the Southern Cross
Roy MacLeod
Royal Society of New South Wales, in association with Sydney University Press
ISBN 9781-9208-9880-9
Wlien Archibald Liversidge first arrived at the
University of Sydney in 1872 as Reader in
Geology and Assistant in the Laboratory, he had
about ten students and two rooms in the main
building. In 1874, he became Professor of
Geology and Mineralogy and by 1879 he had
persuaded the University Senate to open a
Faculty of Science. He became its first Dean in
1882.
In 1880, he visited Europe as a tmstee of the
Australian Museum and his report helped to
establish the Industrial, Technological and
Sanitary Museum which formed the basis of tlie
present Powerhouse Museum’s collection.
Liversidge also played a major role in establishing
the A.ustralasian A.ssociation for the A.dvancement of
Science which held its first congress in 1 888.
This book is essential reading for those
interested in the development of science in
colonial Australia, particularly the fields of
crystallography, mineral chemistry, chemical
geology and strategic minerals pokey.
Archibald
Liversidge
Imperial
Science
under the
Southern
Cross
To order your copy, please complete the Liversidge Book Order Form available at:
http://rovalsoc.org.au/pubHcations/books/McLeod Liversidge Order Form.pdf and return it together
with your payment to:
The Royal Society of NSW,
(Liversidge Book),
PO Box 576,
Crows Nest NSW 1585,
Australia
or contact the Society:
Phone: +61 2 9431 8691
Fax: +61 2 9431 8677
Email: info@royalsoc.org.au
Information for authors
Details of submission guidelines can be found in the on-line Style Guide for Authors at:
http: / / rovalsoc.org.au/ publications / autlior info.htm.
Manuscripts are only accepted in digital format and should be e-mailed to: editor@rovalsoc.org.au
The templates available on the Journal website should be used for preparing manuscripts. Full instructions for preparing
submissions are also ^ven on the website.
If the file-size is too large to email it should be placed on a CD-ROM or other digital media and posted to:
The Honorary Secretary (Editorial),
The Royal Society of New South Wales,
PO Box 576,
Crows Nest, NSW 1585
Australia
Manuscripts will be reviewed by the Editor, in consultation with the Editorial Board, to decide whether the paper will be
considered for publication in the Journal. Manuscripts are subjected to peer review by at least one independent reviewer.
In the event of initial rejection, manuscripts may be sent to other reviewers.
Papers (other than those specially invited by the Editorial Board) will only be considered if the content is either
substantially new material that has not been published previously, or is a review of a major research programme. Papers
presenting aspects of the historical record of research carried out within Australia are particularly encouraged. In the case
of papers presenting new research, the author must certify that the material has not been submitted concurrently elsewhere
nor is likely to be published elsewhere in substantially the same form. In the case of papers reviewing a major research
programme, the author must certify that the material has not been published substantially in the same form elsewhere and
that permission for the Society to publish has been granted by aU copyright holders. Letters to the Editor, Discourses,
Short Notes and Abstracts of Australian PhD theses may also be submitted for publication. Please contact the Editor if
you would like to discuss a possible article for inclusion in the Journal.
The Society does not require authors to transfer the copyright of their manuscript to the Society but authors are required
to grant the Society an unrestricted licence to reproduce in any form manuscripts accepted for publication in the Journal
and Proceedings. Enquiries relating to copyright or reproduction of an article should be directed to the Editor.
Volume 149 Parts 1 & 2 2016
CONTENTS
Numbers 459 to 462
Robert E. Marks: Editorial.
Presidential Address
Donald C. A. Hector. Presidential address.
SMITHSONIAN LIBRARIES
3 9088 01934 7483
1
5
Refereed Papers
William L. Griffin, Sarah E.M. Gain, David T. Adams, Vered Toledo, Norman J. Pearson and Suzanne Y. OReilly: 17
Deep-earth methane and mantle dynamics: insights from northern Israel, southern Tibet and Kamchatka.
Martin G. Banwell, Benoit Bolte, Joshua N Buckler, Ee Ling Chang, Ping Lan, Ehab S. Taher, Lorenzo V White 34
and Anthony C. Willis: Chemoenzymatic pathways for the synthesis of biologically active natural products.
Address
Ian Castles: The curious economist: William Stanley Jevons in Sydney. 5 1
Discourse
Robert E. Marks: William Stanley Jevons, Fellow of the Philosophical Society of N.S.W, 1856-1859. 59
RSNSW Scholarship Winner, 2015
Charles S. P Foster. The evolutionary history of flowering plants. 65
PhD Thesis Abstracts
Jessica Alcorso: Adherence to self-management and psychological distress in women with breast cancer-related 83
lymphoedema.
Nathan Berger. A social psychological examination of factors shaping career and education aspirations through 84
childhood and adolescence.
Nathan Caruana: The cognitive and neural mechanisms of joint attention: a second person approach. 85
Samantha Katherine Dawson: Environmental flows at work; restoring floodplain wetlands through return of 87
historical conditions.
Samantha J. Emery: Quantitative proteomic analyses of isolate variation and virulence in Giardia duodenalis. 88
Benjamin Farr~Wharton: Improving labour outcomes in the creative industries: the role of creative workers’ 90
social network structure and organisational business acumen.
Lynnette Hicks: Dim and dimmer: the production and diffusion of the natural sciences in Australia between the 92
1770s and the 2010s.
Catherine Hoad: Pale Communion: whiteness, masculinity and nationhood in heavy metal scenes in Norway, 94
South Africa and Australia.
David Karel Hutchinson: Interhemispheric asymmetry of global warming: the role of ocean dynamics. 95
Bradley N Jack: Conscious and not-conscious processing of visual mismatch negativity. 96
Todd Jolly: White matter microstructural decline and cognitive performance in older adults: the influence of 97
cardiovascular health.
Gam Thi Hong Lmng. Achieving change in student assessment in Vietnamese teacher training institutions. 98
Brendon Murphy: Zone of impeachment: a post-Foucauldian analysis of controlled operations law and policy. 100
Lucy Murtha: The effects and mechanisms of the therapeutic hypothermia on intracranial pressure regulation 101
following ischaemic stroke in rats.
Patrick Mark Smith: Palaeontology, taxonomy and biostratigraphy of Cambrian assemblages from the 103
Pertaoorrta Group, Amadeus Basin, Northern Territory.
Thi Le Tran: Targeted, one-to-one instruction in whole-number arithmetic: a framework of key elements. 104
Proceedings 106
Awards 108
Information for Authors Inside Back Cover
ISSN 0035-9173
The Royal Society of New South Wales
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Crows Nests NSW 1585^ Australia
info^royalsoc.org.au (general)
editor@royalsoc.org.au (editorial)
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"9
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Published December 2016
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