Annals of Botany 93: 345±351, 2004

doi:10.1093/aob/mch058, available online at www.aob.oupjournals.org

VIEWPOINT

Plant Intelligence: an Alternative Point of View

RICHARD FIRN*
Department of Biology, University of York, PO Box 373, York YO10 5YW, UK

Received: 9 September 2003 Returned for revision: 24 October 2003 Accepted: 8 January 2003

The concept of plant intelligence has been advanced by Trewavas as a potentially useful framework to guide
those seeking to understand plant growth and development. In this short critique, the validity of this concept is
critically assessed. Central to this critique is the proposition that the concept of the individual, to which intelli-
gence and behaviour are intimately linked, cannot usefully be applied to plants. It is argued that the adaptive
responses of plants are best appreciated if the importance of the autonomy of the individual organs is acknow-
ledged. Although Trewavas does acknowledge the autonomy of organs by describing an individual plant as
being `a democratic confederation', that terminology implies a complexity to the interaction between organs
which would demand a cogitative ability beyond that actually demonstrated in plants. It may be more appropri-
ate to consider a plant as operating normally as a simple economic federation of many specialized economies
(organs and cells). Occasionally, there can be a dramatic, and sometimes complex, reshaping of the economic
balances, with the result that the fate of some or many of the individual cells will change. However, such major
changes in growth and development are driven by a few simple events in an individual organ and cells. These
driving events are more akin to small local revolutions in individual states than they are to democratic decisions
in a sophisticated confederation. ã 2004 Annals of Botany Company

Key words: Phenotypic plasticity, adaptation, development, individuality, intelligence, plant behaviour.

INTRODUCTION the overlaying of a very in¯exible pattern of growth and

In a stimulating, thought-provoking and wonderfully wide-
ranging article, Trewavas has argued that `the use of the
term intelligence with regard to plant behaviour will lead to
a better understanding of the complexity of plant signal
transduction, the discrimination and sensitivity with which
plants construct images of their environment and raise
critical questions how plants respond at the whole plant
level' (Trewavas, 2003). Trewavas argued that evidence of
learning and memory in plants, both considered necessary
components of a truly intelligent system, has largely been
ignored with the result that there is little appreciation of
intelligence in plants. While there is a danger that any
discussion about intelligence in plants becomes distracted
by semantics, beneath any such linguistic debate there are
important issues that need to be appreciated by anyone
attempting to understand how plants work.
I N T EL L I G E N C E IS A P RO PE R T Y O F
INDIVIDUALS?
A key issue in discussing intelligence is the concept of
individuality. The word individual leads us to the ®rst
semantic trap. The word in English derives from Latin, in-
meaning not and dividuus meaning divisible. However,
dictionary de®nitions (e.g. Chambers, 1959) stretch this
de®nition to encompass meanings from subsisting as one to
having an independent existence. Animals are highly
individualistic in the sense of subsisting as one. Each
animal has an individual identity which is a consequence of
* For correspondence. Fax 01904 328505, e-mail drf1@york.ac.uk
Annals of Botany 93/4, ã Annals of Botany Company 2004; all rights reserved
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development with an extremely variable behaviour (both
`hard wired' and learned). (This short article focuses on
comparisons of unitary `higher' animals and plants while
recognizing that equally important comparisons might be
drawn between `lower' and colonial animals and plants.) In
contrast, the concept of the individual in the case of a plant
is much harder to de®ne. It is easy to think of a very young
seedling (the commonest experimental object in plant
science) as being an individual, but which is the individual
bracken plant in a huge clonal patch of bracken? Are all
Cox's Orange apple trees the same individual? Maybe,
along with the injudicious use of statistics and the excessive
constraints that tight experimental control impose on studies
of plant functioning (Trewavas, 2003), we need to add that a
blinkered use of `individual' young seedlings as model
systems may also be misleading. Interestingly, the wide-
spread adoption, in the 19th century, of the very young
seedling as the model for much experimental work
coincided with the exploration of the proposal that, as in
animals, certain plant organs could control the growth and
development of other parts of the individual (for some
history, see Went and Thimann, 1937). In a very young
seedling with only two, distinct elongating organs, each
possessing an apical region pioneering the exploration of
their own very different environments, it is maybe reason-
able to consider the seedling as an individual. However,
once the two young axes lose their monopolies when other
organs grow and develop, the sense of what is the individual
becomes less clear. Consequently, while there are some
general principles governing the ways in which plants grow
and develop, principles that may be most easily identi®ed by
 346 Firn Ð Plant Intelligence: an Alternative Point of View
studying very young seedlings, the experimentalist needs to
be conscious of the need to explain how similar processes
might be controlled in more complex, multi-organ plants. A
4-d-old arabidopsis seedling might look like an individual
but a strawberry plant with attached runners, a coppiced
willow tree or a shallot plant present dif®culties for those
seeking to identify the individual. Those experimentalists
who con®ne their studies to young seedlings are not
confronted with one of the key differences between animals
and plants. In animals, growth and development is the
enlargement and modi®cation of the individual. But, in
plants, growth and development also includes the adding of
new members, or the discarding of some old members, of
what Trewavas calls the `democratic confederation'. This
key difference is least apparent in young seedlings; but even
in young seedlings the main organs can be grown in
isolation, hence in the short term such organs largely depend
on their neighbours in the `confederation' as suppliers
(McIntyre, 2001) rather than governors. The `confederation'
has evolved with economic dependence being the main
driving force behind specialization, hence the seedling is
really an economic union rather than democratic confeder-
ation. (Plant biologists might more usefully read Adam
Smith than Thomas Paine to appreciate their experimental
subjects.) The capacity of the seedling, and more evidently
the capacity of the mature plant, is largely the summed
capacity of the component parts, with many of those
capacities being interdependent largely in `economic'
terms. As will now be argued, any `intelligence' that
might be ascribed to `the plant' could only reside in organs,
tissues or cells because the concept of the plant as an
individual is a misleading one.
I N T E LL I G E N C E
`Intelligence is not a term commonly used when plants are
discussed' (Trewavas, 2003). Maybe this is because plants
are clearly not intelligent in the sense that most people
understand the term `intelligent'. The word intelligence
comes from Latin (from intellegere to discern, comprehend
and, literally, `choose between', from inter- and legere, to
choose). The key words in this de®nition are discern,
comprehend and choose, all of which are terms that are
meaningful in the context of human behaviour. These terms,
and the concept of intelligence, were adopted by those
participating in the development of the English language to
describe actions, and to express thoughts, about their own
behaviour. The terms discern, comprehend and choose each
imply a considerable degree of mental processing of more
basic sensory information. There is rather little evidence
that plants (or maybe more accurately plant cells) do
anything other than rudimentary processing of sensed
information and that alone should caution us against
adopting the term `intelligence' when discussing the
abilities of plants. Trewavas moves to ®rmer ground by
adopting Stenhouse's view (Stenhouse, 1974) that intelli-
gence is `adaptive variable behaviour within the lifetime of
an individual', hence directing attention at the term
behaviour. This is convenient for Trewavas's thesis because
many biologists use the term behaviour simply to denote `a
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response to a stimulus' and that allows the term to be
applied rather easily to plants. Plants do indeed show
responses to stimuli and clearly those responses are largely
adaptive, hence one comes to a point that Stenhouse's
de®nition of intelligence could be applied to plants. But has
this semantic exercise helped us identify new, productive
ways of observing, studying, manipulating or understanding
plants? Several generations of scientists have never doubted
that all organisms have evolved to show some ability to
adapt to their environmental circumstances. A bacterium
can monitor its environment and instigate developmental
processes appropriate to the prevailing circumstances, but is
that intelligence? Such simple adaptation behaviour might
be bacterial intelligence but is clearly not animal intelli-
gence. Hence we have reached a de®nition of intelligence
that has no meaning unless combined with another word or
used in a precise context in a sentence. This is not a new
problem, as shown by the debate about whether machines
can be intelligent. The term machine intelligence can mean
something other than the intelligence of a machine. We can
certainly start using the term plant intelligence but only if it
is agreed that the term has nothing to do with intelligence
(the ability to discern, comprehend and choose) in the more
widely accepted sense. (To add to the opportunity for
confusion it should be pointed out that the term `plant
intelligence' is already in use to mean the utilization of
data about the performance of factories to optimize
performanceÐbut plants could be considered as factories,
so maybe that is OK; http://www.gefanucautomation.com/
plantintelligence/default.asp.) However, why do we need a
new term? If the discussion of a plant's adaptive responses
is less ambiguous than a discussion of the plant's intelli-
gence, why adopt the new term? Indeed, might not the
adoption of the term plant intelligence begin to distort,
rather than clarify, our perception of the way in which plants
function? To illustrate this danger, some of the key terms
that are part of Trewavas's view of plant intelligence will
now be placed under scrutiny.
LEARNING
Trewavas (1999, 2003) considers learning to require two
faculties. The organism must be capable of having a goal,
and the organism must be able to assess its current
behaviour and adjust that behaviour to make the attainment
of the goal more likely. As part of the latter ability, he
considers the need for an error-detection mechanismÐa
method for judging current state against a reference state
(goal). Each of these elements needs to be considered from
the perspective of the adaptive responses of plants.
Goals
Trewavas de®nes a goal as a plant's ®tness objective. The
main problem with the concept of a single plant possessing a
goal is that each mature plant is made up of ever-changing
components (roots, leaves, ¯owers, etc.), each of which
occupy their own temporally and spatially variable environ-
ments. When a plant is at its most individualistic, as a seed,
there are clear goalsÐdispersal, survival and germination at
 Firn Ð Plant Intelligence: an Alternative Point of View
the right time. However, shortly after germination each
extending meristematic region, such as a root tip or an apical
bud, may encounter local conditions that are very different
from those encountered by other organs on the same plant
exploring other localities. Furthermore, the conditions
experienced by one generation of organs are likely to differ
from those experienced by later generations. The need to
optimize the performance of each organ in its own variable,
unpredictable environment has given rise to a selective
process that has favoured specimens with sensing and
response capacities, which are largely devolved to the organ
or sub-organ level. This devolution of sensing and response
to functionally specialized organs results in most goals
being themselves functional and local. Thus it is hardly
surprising that the great majority of sensing and response
events in plants are localized to the cell, tissue or organ, in
that order. Furthermore, because most organs are multi-
functional and grow in heterogeneous environments, any
overall goals must be ¯exible and necessarily based on a
conservative compromise. To cope with the variability that
an organ may encounter, it is likely that evolution will have
selected for variants that can make approximately the
optimum number of structures (be they organs, cells or cell
components), but where there is some variable functional
capacity. The variable functional capacity of each unit
enables the overall current and short-term functional needs
to be met. For example, the gas exchange capacity of a leaf
is in part determined by the number and the aperture of the
stomata, and varying the latter is the most effective way of
giving operational control to the leaf in a varied environ-
ment. Furthermore, the environmental conditions that a
developing leaf experiences might differ considerably from
the conditions that a mature leaf experiences. So for an
organ there may be one or more developmental goals and
some operational goals. The end result should provide
suf®cient functional ¯exibility to cope ef®ciently with
whatever changing environment it may have to operate in.
Another level of control might have evolved to cope with
rarer, more extreme conditions that push organs beyond the
limits of their operational controls. Such `emergency'
shutdown routines at best protect all the members of the
economic union and at worst ensure the survival of
suf®cient members to enable a successful recovery
programme to be initiated. The graded responses of plants
to increasingly severe water stress would be an example of
such `emergency' recovery routines (Hsiao, 1973). Even in
these circumstances, the response of `the plant' is actually
the sum of the responses that take place in cells, tissues and
organs. In such circumstances, each responding cell need
have no information as to the location or scale of the
problem elsewhere, simply that a programmed response is
needed to contribute to the ®tness of the union.
In conclusion to this consideration as to whether a plant
can possess a goal, the most important means by which the
economic union that is the plant achieves the fundamental
goal common to all living creatures (survivorship and
reproduction) is to possess both a ¯exible way of controlling
the number and types of organs (organs can be initiated and
discarded) and allowing each organ considerable oper-
ational autonomy. Most of the developmental and oper-
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347
ational control of the organs is devolved to local sensing, so
that the development and functioning of an organ can match
the local conditions and hence can contribute most effect-
ively to the union. Selection in plants has favoured the
evolution of ¯exible modular organisms where the ®nal
structure is just one of many different possible outcomes.
The developmental ¯exibility which enhances plant ®tness
is in contrast to the adaptive ¯exibility that behaviour gives
to animals.
Error detection
As outlined in Box 1, a simple on/off switch linked to a
sensor is all that is needed for `error detection'. The
examples of oscillations that Trewavas takes as evidence of
`trial-and-error learning' could equally be used as evidence
for simple automaton behaviour. Examples of homeostatic
mechanisms in plants and animals, which are readily
modelled in terms of a sensor and a reaction to sensor
state, provide robust, if simple control systems, suf®cient for
many purposes. The fact that many examples of oscillatory
behaviour occur in ephemeral organs also causes one to
doubt whether any bene®t would accrue from the possession
of an ability to learn. For example, a root undergoing
gravitropism is actually a different root at the start and end
of the process. If a root is exposed to an extreme
gravitational stimulus twice, 12 h apart (which would be
an event so rare in nature that one cannot imagine selection
optimizing a mechanism that included a learning stage), the
second stimulus will initiate a new response in cells that
have not necessarily experienced the ®rst stimulation, and
hence they cannot usefully have learned. This is in contrast
to an animal where learned behaviour resides at the
organism level ± the hand has no memory of being burned,
but the brain controlling the hand might possess the
appropriate knowledge after an adequate previous learning
experience.
Choice
Trewavas considers the evidence that plants make
choices, or decisions, in reaching his conclusion that plants
have a kind of intelligence. But can plants really make
choices in any meaningful way? A thermostat can `make a
decision' (Box 1) but few would claim that it makes a real
decision in any sense of free will. There would seem to be a
confusion here between approaching a state with a limited
number (often two) of predetermined outcomes and real
decision making. A more demanding criterion for choice or
decision making is whether there are several variable
outcomes, e.g. action or inaction, and different ways of
taking action. When `choices' are made in plants they are
made at the organ (or cell) level and not at the plant level.
Even for the most fundamental `choice' that `a plant' might
makeÐto initiate ¯oweringÐone could argue that the
switch to reproductive development in the apex is a simple
two-state system. It is also surely signi®cant that the control
of ¯owering is not determined centrally by `the plant',
indeed the photoperiodic sensing can occur in many leaves
or even within a small part of one leafÐa very robust
 348 Firn Ð Plant Intelligence: an Alternative Point of View
system (Zeevaart, 1976). The plant does not make a choice
to ¯ower, instead a leaf in¯uences the predetermined fate of
some cells in the apex. There is no choice made by `the
plant' rather events occur within the plant that dramatically
change the balance within the economic union, with
consequent changes to the fate of some members of the
union.
SPATIAL MAPS
Does a plant build up a spatial map of its environment? To
answer this question one needs to think more carefully about
the type of plant, the type of environment and type of spatial
map that might provide a ®tness bene®t. Once again, the
favoured experimental subject of those investigating a
sensing system in plants, the very young seedling grown in a
very controlled environment, might be misleadingly simple.
It is very clear that the construction and maintenance of an
accurate spatial map of a large plant such as a tree would
demand huge processing power. Thousands of leaves and
roots, each having a need to sense their local environment to
serve their own needs, would produce very large amounts of
data. The data would be changing by the minute as the
leaves ¯uttered in a breeze and the roots explored their very
heterogeneous environment. This example indicates that
there is a need for great caution when scaling up knowledge
from young seedlings to more mature, more complex plants.
It might be more pro®table to regard a tree not as a big
seedling but as a collection of thousands of seedlingsÐeach
potential cutting is a potential new plant, hence the concept
has some validity. This consideration should make us
concentrate on the organizational level at which spatial
mapping could be meaningfulÐthe organ level. In the main,
each operational or developing organ can, at best, sense a
few parameters that suf®ciently characterize their local
environment with respect to the function of that organ. Each
organ has some extra information derived from the overall
integrative state of the plant (much of which might be
considered to be similar in nature to consequences of
independent, competing economic activities in a human
confederation). This extra information is available from the
summed activities of all members of the federation of
organs, but such exotic information is rarely complex. Some
extra speci®c information might also be available to certain
cells from chemical signals emanating from non-adjacent
cells. Although Trewavas, as Canny (1985) before him,
argues that the pathways of inter-organ transport could
allow complex chemical signals to ¯ow between organs, it is
unlikely that the complexity of the signalling molecules
implies a complexity of message. (It should also be noted
that the existence of signalling systems is not the issue at
stake, rather it is whether complex signalling is a good
indicator of the existence of an `intelligent' adaptive
response.) The complexity of signalling molecules is only
necessary to ensure that the right recipient receives the
simple message. A signalling molecule is akin to a postcard
bearing a tick or a cross as the message, with the much more
complex information of the address ensuring that the simple
message reaches the right recipient (Firn, 1985). [The reader
interested in the complexities of hormonal signalling in
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plants is referred to Weyers and Paterson (2001).] It might
also be signi®cant that some of the best examples of long-
distance signalling (e.g. ¯owering or tuberization) are
actually ones where spatial information is not importantÐ
many leaves can detect day-length changes and it matters
not which one does. Are there any examples of spatial
information derived by plants that is not used mainly
locally?
In conclusion, there is no convincing evidence that plants
possess the ability to convey suf®cient information from
sensors to any processing units to enable a meaningful
spatial map to be constructed. The only map that a plant as a
whole makes of its environment is recorded in its morph-
ology, anatomy and chemistry. It is a map of past events, not
of future plans.
M EM O R Y
There can be no doubt that the developmental history of a
plant can determine the outcome of many subsequent
sensing events. More accurately, the developmental history
of many organs (or cells) can determine the response that
results from subsequent stimulation. Is that memory or is it
simply a consequence of developmental progression? A
simple washing machine controller possesses `memory' due
to the equivalent process of developmental progression.
Developmental progression at its simplest is a linear
sequence of events, with one or more steps under the
control of some sensor (Box 1). Once the sequence is
initiated, progress is determined by the state of some or all
of the controlling sensors. Events occurring at an early stage
can in¯uence the outcome of later events and the system can
thus have `a memory' of earlier events. However, a crucial
distinguishing characteristic of memory in animals is that
the processes are not linear and they are not sequence-
dependent.
Assuming that `plant memory' is little more than
developmental progression, it is clear once again that
developmental progression would be a property of organs or
cells and not plants. This fact reveals why it is unlikely that
true memory could have evolved in plants. Organs, for
example, need to cope with the environment in which they
®nd themselves and, being essentially ephemeral, they
could bene®t little by retaining and using information of
past events. A root growing in a heterogeneous environment
cannot use an experience of its current state to predict its
future state. Water (or potassium or nitrate, etc.) which is in
short supply now may be in ample supply later. The memory
of animals is aimed at using the experience of one organ to
the advantage of othersÐa hand that is burned can provide a
lesson to the whole organism so that in future other limbs
can avoid experiencing the same problem. Where is the
equivalent collective organism experience (= memory) of
bene®t in a plant's growth and development?
CONCLUSION
Adaptive behaviour takes many forms in different organ-
isms. By using terms and concepts of learning, memory and
intelligence that derive from human experience one can
 Firn Ð Plant Intelligence: an Alternative Point of View
build up a view of each type of adaptive behaviour in other
organisms, but maybe only at the expense of clarity. Indeed,
there is a danger that the adoption of concepts and terms
from higher animals distracts attention away from the key
adaptive processes evolved by organisms that have not been
selected to learn, memorize and think. In this article it has
been argued that the `intelligence' perceived in the adaptive
response of a plant might best be considered to be the sum of
the collective adaptive responses of its cells. A general
theme running through this article is that the human sense of
the individual easily distorts the ability of humans to
appreciate how a plant operates. Even in the previous
sentence the two simple words `a plant' feel, to the author,
inappropriate because these words emphasize the individual
entity. Plant population biologists have adopted terms such
as genet and ramet (Harper, 1977) because they appreciate
that descriptors beyond the concept of the individual are
needed to understand plant populations. However, many
plant biologists studying the functioning of cells or organs
don't look to plant population biologists for inspiration but
to animal biologists. Such plant biologists might bene®t
from shifting their focus to other types of plants rather than
other types of organisms. An appreciation may then develop
that clonal plants are simply demonstrating an extreme form
of developmental adaptation, but it is a form of adaptation
that can be seen in simpler terms in a 4-d-old arabidopsis
seedling. In clonal plants, the ramet is a device to provide an
ef®cient division of labour in a heterogenious, patchy
environment (Stuefer et al., 1996; Hutchings and
Wijesinghe, 1997; JoÂnsdoÂttir and Watson, 1997) and that
is simply an extension of the strategy of allowing organs an
autonomy to optimize their localized performance. The
extent to which those semi-independent units interact, and
how they interact, is easier to understand if the overall
strategy of plant development is comprehended.
Finally, the author, during the preparation of this article,
has become very aware that human language guides and
greatly limits our thoughts when trying to appreciate the
functioning of plants. It might ultimately be unproductive to
debate whether a plant has memory, whether a plant can
learn or whether a plant can possess a spatial map, because
the key terms come from an organism that is highly
individual, an organism that ®nds it hard to appreciate the
faculties of organisms that lack individuality and all that that
implies. Our language lacks appropriate words. However, if
new words are needed to describe how plants function,
maybe we should invent new ones rather than trying to
rede®ne existing ones.
ACKNOWLEDGEMENTS
Tony Trewavas is thanked for many years of provocative
and stimulating comment. John Raven is thanked for
sharing his wisdom and knowledge.
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349
BO X 1
Intelligence: switches, transistors, logic gates and
computers
How much complexity must a system possess before
it is regarded as intelligent? The way in which
relatively simple machines can be built to carry out
quite complex tasks can help us answer this question.
In a simple refrigerator, a temperature-controlled
switch (thermostat) simply turns the cooling system
on when there is deviation from a prede®ned
temperature. The refrigerator has been set a goal and
achieves it by what could be considered to be error
detection. However, such a simple control system is
very prone to the kinds of oscillations that impress
Trewavas. Due to the thermal mass of the thermostat
and the thermal mass of the cooling coils, once the
thermostat removes the power from the compressor,
cooling continues beyond the value set by the
thermostat. So the oscillations that Trewavas con-
siders evidence of sophisticated `learning' can also be
characteristic of simple on/off controls, lacking
hysteresis. More sophisticated controls can eliminate
temperature oscillations in a refrigerator. A controller
(known as a proportional controller) that can measure
the magnitude of deviation from a set point, and can
modulate the magnitude or duration of the restoring
process, will be able to control a process with much
less oscillation. In plants, where the control mechan-
isms are often devolved to the cell level and where the
behaviour of the cells may not be synchronous, it is
possible that simple control systems suf®ce and that
oscillatory behaviour is to some extent mitigated by
the hysteresis capacity given to the system by the fact
that multiple sensors/regulators are aggregated at the
organ level.
By combining a few simple sensors with a timing
mechanism, more sophisticated, but still simple,
control systems can be built. A well-known example
is the domestic automatic washing machine. These
appliances have sometimes been given names or
descriptions by their manufacturers that imply an
intelligence. However, in most washing machines, the
`intelligence' actually resides in the user and the
designer of the controller. The user selects a goal
(from a choice of goals actually determined by the
design engineer) and the controller is designed to start
and stop certain processes in order to progress towards
the chosen goal. To achieve this, several concentric,
plastic pro®les are rotated by a motor and the peaks
and troughs on each pro®le activate one on/off switch.
Each switch controls one of the main power circuitsÐ
the main drum motor, the draining pump, the water
¯ow valves, the timer motor and the heater. A few
simple on/off sensors (water temperature and water
depth) interact with the motor that drives the spindle
on which the pro®les rotate. In other words, complex
temporal sequences, tied into several physical param-
eters, can be achieved by no more than a few on/off
 350 Firn Ð Plant Intelligence: an Alternative Point of View
switches linked to some timing mechanism. Such a
mechanical timer even has `memory' in that if its
cycle is interrupted, it starts again where it has
stopped. It also has `memory' in terms of being able to
proceed to one process only after another, prede®ned,
process has been ®nished (e.g. it won't spin your
clothes until the drum is free of water). Thus a modern
washing machine controller could be analogous to
developmental progression in a plant ± processes
happen in a sequence and there are sensors that are
made active for de®ned periods and which can
in¯uence the rate of progression itself.
The machine analogies becomes more interesting if
one considers what happened after the transistor was
invented ± the evolution of complex mechanical
controls eventually ceased. Mechanical controllers
persisted for decades (indeed, they are still used in
some washing machines) because they were reliable,
robust and cheap. However, the invention of the
simple transistor opened up an evolutionary path that
would lead to controllers with more precision, more
versatility and lower costs. The transistor can be
regarded as an electrical on/off switch. Within 10
years of its invention it had become possible to
produce devices that incorporated several transistors
as cheaply as producing one transistor. By connecting
several on/off transistor switches, such that speci®c
inputs and outputs were connected, it was possible to
produce electrical circuits that displayed Boolean
logic. The development of logic circuits which
retained information (memory) and which could
measure time, enabled the development of machines
that could monitor many inputs, process the informa-
tion in complex ways and produce a variable output
dependent on the outcome of the result of the
processing of that information ± the simple computer.
The myriad of electronic devices currently available is
simply the result of an evolutionary process in an
industry that has had to learn new ways of connecting
up transistors in order to survive.
So where are bacteria, plants, simple animals and
humans on the continuum of machine intelligence that
stretches from the thermostat to the computer?
Bacteria would seem to be at the simple end of the
spectrum (cheap washing machine). At the other
extreme, the evolution of the nervous system in
animals could be regarded as the equivalent of the
evolution of industrial control processes after the
invention of the transistor. In the same way that
linking up transistors in various ways gave rise to
memory and machine learning, so the linking of cells
in the nervous system enabled analogous systems to
evolve in animals. The nervous system of an animal
has a `hard-wired' capacity to ensure that basic
physiological functioning operates optimally (a kind
of collective species experience) but it is also
programmable (what we call learned behaviour) to
enable each individual to gain and use their own
experience to adapt to their own circumstances. This
programmable element is the capacity for true learn-
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ing that is characteristic of animals and it is a
characteristic that distinguishes animals from plants.
Where are plants on this continuum of control
process sophistication? As argued elsewhere in this
article, plants are modular organisms with some
information ¯ow between organs. Quite sophisticated
control systems can be elaborated by using a hierarchy
of controls ± some local and some more regional.
Consider the operation of a heating system in an
apartment block with a central hot water heating
system. The control for the temperature for most
rooms is devolved to the user of that room (via a crude
thermostatic radiator valve) but each apartment can
control when heating should be available (time switch
controlling an on/off diverter valve connecting the
main heating supply pipe to the apartment's heating
system). The apartment occupier can also set a lower
minimum level ± a simple thermostatic switch that
bypasses the timer. The source of overall heat for the
apartment block is controlled by sensors on the ¯ow
and return pipes and the output of the heat source is
proportionally regulated to meet demand. To antici-
pate demand somewhat, sensors for the outside
temperature are also built in to the control system of
the central heat source. In other words a devolved,
hierarchical system of simple controls meeting local
needs, overlaid by a few layers of controls operating at
a higher level, largely by monitoring the state of major
¯ows into and out of the system, can provide an
adequately regulated, robust and versatile system
which would work in most environments. If you add
to such a hierarchical system some speci®c opportun-
ities for the communication between some local
control systems, then even more complexity is
possible. The modern alternative to such a system of
linked, local, hierarchical controls (which may be
what is found in plants) would be a central controller
with a capacity to ®ne tune each control element via
learning and memory (which would seem to be
analogous with animal control systems).
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