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Smells like hive spirit

The Spirit of the Hive - The Mechanisms of Social Evolution by Robert E. Page, Jr., Harvard University Press, 2013.
US$39.95/29.95/s36.00, hbk (xiv + 226 pp.), ISBN 978-0-674-07302-9
Ulrich R. Ernst
Research Group of Functional Genomics and Proteomics & Laboratory of Socioecology and Social Evolution, Department of
Biology, KU Leuven, Naamsestraat 59, bus 2465, 3000 Leuven, Belgium
There is an abundant offer of academic
books on the topic of social insects
currently available, for example [13].
Among these, the new book by Robert E.
Page, titled The Spirit of the Hive, stands
out in its analysis of social evolution due to
its integration of seemingly disparate
topics, such as foraging preferences, sen-
sitivity to sugar, evolution of multiple
mating, and division of labour with task
specialisation. Page is not asking the traditional ecological
questions (for instance, kin selection is only mentioned
once simply to announce that it will not be discussed at all
in this volume), nor does he use an overly theoretical
approach. Rather, he chooses to examine the inner working
of a colony and the mechanisms that govern its organisa-
tion by studying the causes of individual behaviour.
He focuses on the honeybee, the most studied social
insect, to investigate how tens of thousands of individuals
in a colony can cooperate and know when to perform
specic tasks to create a functioning society within the
hive. Because the bees are not given orders by the queen,
one must assume that there is another inuence organis-
ing the workforce of a hive. This is what the Belgian Nobel
prize winner for literature, Maurice Maeterlinck, imagined
in 1901 as the spirit of the hive [4]. In his book, Page
demonstrates that genetics and physiology can explain
what poetry left to idealistic reverie.
Within ten chapters, Page develops an increasingly
complex model, the Reproductive Groundplan Hypothesis
[5], that can account for most phenomena related to task
specialisation, individual reproductive and foraging deci-
sions, and colony level traits. He keeps the descriptions and
explanations simple and covers basic biological principles
when necessary (e.g., when talking about RNA interfer-
ence experiments, he briey reviews DNA, RNA, and
proteins). In the same vein, Page follows the advice of
his mentor, Harry Laidlaw, by using redundancies to make
his points clear. Each chapter is illustrated with tables,
photographs, diagrams, cartoons, and schemes throughout
to help the reader understand the hypotheses and experi-
mental data that lead to the model. Occasionally, however,
some of the graphs are too schematic to convey the
intended information. For example, an ellipse containing
three circles and little ladders might evoke associations of
a cell with multiple nuclei, especially because the legend
explains how DNA is translated into RNA in the nucleus.
Yet, this was meant to symbolise the working of RNA
interference in the fat body of a bees abdomen.
This volume is not intended for laymen. Yet, thanks to
Pages efforts to make the story of the genetic architecture of
a social insect colonyaccessible to a broadaudience, theycan
still grasp the gist, even though they might not be able to
fully appreciate all the arguments presented in this book.
However, for biologists, the ndings from behavioural
experiments, colony level traits, selective breeding and
crossing, dissections, mathematical models, and genetic
mapping are clearly presented, the results thoroughly dis-
cussed, and the consequential experiments sufciently mo-
tivated. Indeed, this is the strength of this thorough yet
unpretentious work: combining seemingly eclectic data,
gathered by a wide array of methods, presenting them in
a logical order, and tting them in the unifying hypothesis
represented by the Reproductive Groundplan Hypothesis.
Page generally succeeds in reducing complicated hypothe-
ses and experimental designs to simple questions, as well as
distilling the essence of eachpiece of evidence. By answering
these questions step by step, we get increasingly closer to a
detailed picture of how selection has acted on the genetic
material of solitary ancestors to bring about intricate socie-
ties. Page presents the steps that have been taken in such a
well thought out way that the reader can often anticipate
which experiment will come next.
Even though the evidence for the Reproductive Ground-
plan Hypothesis is convincing, Page rightly warns that our
models are probably (partially) wrong and may be over-
thrown one day. He is especially sceptical about beanbag
thinking and advocates an ecology of genes, in which the
whole genome and the physiology are integrated.
His profound knowledge of the literature, not only of
often widely scattered, published manuscripts but also of
unpublished data, puts him in an ideal position to summa-
rise and synthesise over 30 years of research. In this light,
it is unfortunate that Page decided not to cite the relevant
references in the text, and instead chose to provide a list of
suggested reading for each chapter. Thereby, several arti-
cles are recommended repeatedly in different chapters,
whereas others go unreferenced, even though the authors
are mentioned in the text. However, large portions of the
nal ve chapters of the book can also be found in a recent
review article [6], and its reference list could serve as a
starting point into the literature.
The Spirit of the Hive is a warm invitation and excellent
opportunity for those biologists not familiar with the phys-
iology and genetics of social insect colonies to be introduced
to proximate aspects of social evolution. Readers will nd a
Book Reviews
Corresponding author: Ernst, U.R. (Uli.Ernst@bio.kuleuven.be).
686
seamless tale: genetic changes in physiology inuence
behaviour resulting in different colony level traits altering
the environment of a colony that again impacts physiology
and genetics.
References
1 Holldobler, B. and Wilson, E.O. (2009) The Superorganism, W.W.
Norton, (New York and London)
2 Seeley, T.D. (2010) Honeybee Democracy, Princeton University Press,
(Princeton and Oxford)
3 Turillazzi, S. (2012) The Biology of Hover Wasps, Springer, (New York)
4 Maeterlinck, M. (1901) The Life of the Bee, Dodd, Mead &Co, (NewYork)
5 West-Eberhard, M.J. (1996) Wasp societies as microcosms for the study
of development and evolution. In Natural History and Evolution of
Paper-wasps. (Turillazzi, S. and West-Eberhard, M.J., eds), pp. 290
317, Oxford and New York, Oxford University Press
6 Page, R.E. et al. (2012) Genetics of reproduction and regulation of
honeybee (Apis mellifera L.) social behavior. Annu. Rev. Genet. 46, 97119
0169-5347/$ see front matter
http://dx.doi.org/10.1016/j.tree.2013.09.010 Trends in Ecology & Evolution,
December 2013, Vol. 28, No. 12
How to view our/the universe
The Science of Discworld IV: Judgement Day by Terry Pratchett, Ian Stewart, and Jack Cohen. Ebury Press, 2013. 18.99, hbk (342 pages),
ISBN: 978 0 09 194979 2
Ian C.W. Hardy
School of Biosciences, University of Nottingham, Sutton Bonington Campus, LE12 5RD, UK
Three previous books by the same trio of
authors have explored a multitude of sci-
entic topics, including plenty of biology,
using chapters of real-science inter-
spersed with illustrative and enjoyable
nonscience from the ctional unreality of
Discworld [1,2]. There is now, after an 8-
year gap, a fourth part to the trilogy.
Judgement Day follows the same inter-
spersed structure, but the science:ction
ratio is noticeably higher: the pure-Discworld chapters are
short.
The theme running throughout the book is that there
are two fundamentally different ways to think about the
world. Perhaps not super-surprisingly, these two ways are
essentially science and faith or, more informatively put,
seeing the universe as the context for humanity versus
seeing the humanity as the context for the universe. If we
take the human-centred stance, this is our universe, with
us as its purpose and at the (conceptual if not actual)
centre of it. Then, among other things, the universe is a
resource for us to exploit. If the universe-centred stance is
taken, then the universe is indifferent to our existence and
we are minor players on cosmic scales of time and space.
The value of the latter stance is that it steps away from the
innate human tendency to rely on intuition: scientists
actively try to disprove things that they would like to be
true and build an analytically based understanding of
their surroundings.
The authors claim that thinking without what they term
adequate equipment is often signalled in popular debates
by the phrase I reckon. They are not theologians (although
Cohen was once to become a Rabbi), and neither am I, so I
wonder what our brethren in the humanities will make of
Pratchett et al.s reckoning that few faith-based systems
advocate self-doubt as a desirable instrument of change
and the associated assertion that In religion, doubt is often
anathema: what counts is how strongly you believe things.
I suspect that theologians test their faith more than is
suggested here, at least for a given value of test.
Overall, Pratchett et al. take us engagingly and, at times
brilliantly, through a slew of scientic and mathematical
subjects, using sources from the ancient to the bang-up-to-
date. For readers of TREE, the list notably includes: Dar-
winism, RNA, the importance of ribosomes, astrobiology,
Bayesian neuroscience, and a memorable fridge-magnet-
type phrase Biology isnt just physics and chemistry with
knobs on. In fact, by reading this book, one usefully
improves and consolidates ones grasp of things such as very
small physics (e.g., fundamental particles: in a nutshell,
there seem to be about 17) and very big physics (e.g., the
shape of the universe: it is unknown) more than one does of
the life sciences.
Despite it not containing all that many pages explicitly
devoted to ecology or evolution, I would recommend this
book to readers of TREE for two reasons. Generally, be-
cause thinking about the philosophical method of science is
always useful. Specically, because, like it or not, our
discipline is connected to issues of faith [3], for example,
via debates about intelligent design and religious objec-
tions to genes being selsh. This book provides a relatively
conciliatory discussion, appreciative of some aspects of
faith-based thinking and lacking (most of) the beagle-bite
of Dawkinss God Delusion [4]. When reading near the
beginning, I thought the authors might end up positing
something akin to Goulds non-overlapping magisteria [5].
In fact, the book progressively becomes clear in its uni-
verse-centred, and denitely antifundamentalist, stance.
Judgement Day would nestle happily next to Dawkins [4]
in the library of the Unseen University in Discworld and it
should certainly be placed in the science rather than the
fantasy section (so, not next to the Bible, as nally proposed
in Pratchett et al.s Epilogue). Judgement Day will be of
thought-provoking relevance to ecologists and evolutionary
biologists for as long as there remain two fundamentally Corresponding author: Hardy, I.C.W. (ian.hardy@nottingham.ac.uk).
Book Reviews Trends in Ecology & Evolution December 2013, Vol. 28, No. 12
687

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