Nobel laureate Erwin
ShrÃ¶dinger's What is Life? is one of the great
science classics of the twentieth century. A distinguished
physicist's exploration of the question which lies at the
heart of biology, it was written for the layman, but proved one
of the spurs to the birth of molecular biology and the subsequent
discovery of the structure of DNA. The philosopher Karl Popper
hailed it as a 'beautiful and important book'. It appears
here together with Mind and Matter, his essay investigating a
relationship which has eluded and puzzled philosophers since the
earliest times. Brought together with these two classics are
ShrÃ¶dinger's autobiographical sketches,
published and translated here for the first time, which offer a
fascinating fragmentary account of his life as a background to
his scientific writings.
'This book is a gem with many facets.., one can read it in a few hours; one will not forget it in a lifetime.' Scientific American
'Erwin ShrÃ¶dinger, iconoclastic physicist, stood at the pivotal point of history when physics was the midwife of the new science of molecular biology. In these little books he set down, clearly and concisely, most of the great conceptual issues that confront the scientist who would attempt to unravel the mysteries of life. This combined volume should be compulsory reading for all students who are seriously concerned with truly deep issues of science.' Paul Davies
Order, Disorder and Entropy
Nec corpus mentem ad cogitandum, nec mens corpus ad
motum, neque ad quietem, nec ad aliquid (si quid est) aliud
determinare potest.* SPINOZA, Ethics, Pt In, Prop.2
(Neither can the body determine the mind to think, nor the mind determine the body to motion or rest or anything else -- if such there be.)
A REMARKABLE GENERAL CONCLUSION FROM THE MODEL
Let me refer to the phrase on p. 62, in which I tried to explain that the molecular picture of the gene made it at least conceivable that the miniature code should be in one-to-one correspondence with a highly complicated and specified plan of development and should somehow contain the means of putting it into operation. Very well then, but how does it do this? How are we going to turn ‘conceivability’ into true understanding?
DelbrÃ¼ck's molecular model, in its complete generality, seems to contain no hint as to how the hereditary substance works. Indeed, I do not expect that any detailed information on this question is likely to come from physics in the near future. The advance is proceeding and will, I am sure, continue to do so, from biochemistry under the guidance of physiology and genetics.
No detailed information about the functioning of the genetical mechanism can emerge from a description of its structure so general as has been given above. That is obvious. But, strangely enough, there is just one general conclusion to be obtained from it, and that, I confess, was my only motive for writing this book.
From DelbrÃ¼ck's general picture of the hereditary substance it emerges that living matter, while not eluding the ‘laws of physics’ as established up to date, is likely to involve ‘other laws of physics’ hitherto unknown, which, however, once they have been revealed, will form just as integral a part of this science as the former.
ORDER BASED ON ORDER
This is a rather subtle line of thought, open to misconception in more than one respect. All the remaining pages are concerned with making it clear. A preliminary insight, rough but not altogether erroneous, may be found in the following considerations:
It has been explained in chapter 1 that the laws of physics, as we know them, are statistical laws.* They have a lot to do with the natural tendency of things to go over into disorder.
But, to reconcile the high durability of the hereditary substance with its minute size, we had to evade the tendency to disorder by 'inventing the molecule', in fact, an unusually large molecule which has to be a masterpiece of highly differentiated order, safeguarded by the conjuring rod of quantum theory. The laws of chance are not invalidated by this 'invention', but their outcome is modified. The physicist is familiar with the fact that the classical laws of physics are modified by quantum theory, especially at low temperature. There are many instances of this. Life seems to be one of them, a particularly striking one. Life seems to be orderly and lawful behaviour of matter, not based exclusively on its tendency to go over from order to disorder, but based partly on existing order that is kept up.
To the physicist - but only to him - I could hope to make my view clearer by saying: The living organism seems to be a macroscopic system which in part of its behaviour approaches to that purely mechanical (as contrasted with thermodynamical) conduct to which all systems tend, as the temperature approaches the absolute zero and the molecular disorder is removed.
The non-physicist finds it hard to believe that really the ordinary laws of physics, which he regards as the prototype of inviolable precision, should be based on the statistical tendency of matter to go over into disorder. I have given examples in chapter 1. The general principle involved is the famous Second Law of Thermodynamics (entropy principle) and its equally famous statistical foundation. On pp. 69-74 I will try to sketch the bearing of the entropy principle on the large-scale behaviour of a living organism - forgetting at the moment all that is known about chromosomes, inheritance, and so on.
LIVING MATTER EVADES THE DECAY TO EQUILIBRIUM
What is the characteristic feature of life? When is a piece of matter said to be alive? When it goes on 'doing something', moving, exchanging material with its environment, and so forth, and that for a much longer period than we would expect an inanimate piece of matter to 'keep going' under similar circumstances. When a system that is not alive is isolated or placed in a uniform environment, all motion usually comes to a standstill very soon as a result of various kinds of friction; differences of electric or chemical potential are equalized, substances which tend to form a chemical compound do so, temperature becomes uniform by heat conduction. After that the whole system fades away into a dead, inert lump of matter. A permanent state is reached, in which no observable events occur. The physicist calls this the state of thermodynamical equilibrium, or of 'maximum entropy'.
Practically, a state of this kind is usually reached very rapidly. Theoretically, it is very often not yet an absolute equilibrium, not yet the true maximum of entropy. But then the final approach to equilibrium is very slow. It could take anything between hours, years, centuries,... To give an example - one in which the approach is still fairly rapid: if a glass filled with pure water and a second one filled with sugared water are placed together in a hermetically closed case at constant temperature, it appears at first that nothing happens, and the impression of complete equilibrium is created. But after a day or so it is noticed that the pure water, owing to its higher vapour pressure, slowly evaporates and condenses on the solution. The latter overflows. Only after the pure water has totally evaporated has the sugar reached its aim of being equally distributed among all the liquid water available.
These ultimate slow approaches to equilibrium could never be mistaken for life, and we may disregard them here. I have referred to them in order to clear myself of a charge of inaccuracy.
IT FEEDS ON 'NEGATIVE ENTROPY'
It is by avoiding the rapid decay into the inert state of 'equilibrium' that an organism appears so enigmatic; so much so, that from the earliest times of human thought some special non-physical or supernatural force (vis viva, entelechy) was claimed to be operative in the organism, and in some quarters is still claimed.
How does the living organism avoid decay? The obvious answer is: By eating, drinking, breathing and (in the case of plants) assimilating. The technical term is metabolism. The Greek word [omitted] means change or exchange. Exchange of what? Originally the underlying idea is, no doubt, exchange of material. (E.g. the German for metabolism is Stoffwechsel.) That the exchange of material should be the essential thing is absurd. Any atom of nitrogen, oxygen, sulphur, etc., is as good as any other of its kind; what could be gained by exchanging them? For a while in the past our curiosity was silenced by being told that we feed upon energy. In some very advanced country (I don't remember whether it was Germany or the U.S.A. or both) you could find menu cards in restaurants indicating, in addition to the price, the energy content of every dish. Needless to say, taken literally, this is just as absurd. For an adult organism the energy content is as stationary as the material content. Since, surely, any calorie is worth as much as any other calorie, one cannot see how a mere exchange could help.
What then is that precious something contained in our food which keeps us from death? That is easily answered. Every process, event, happening - call it what you will; in a word, everything that is going on in Nature means an increase of the entropy of the part of the world where it is going on. Thus a living organism continually increases its entropy - or, as you may say, produces positive entropy - and thus tends to approach the dangerous state of maximum entropy, which is death. It can only keep aloof from it, i.e. alive, by continually drawing from its environment negative entropy - which is something very positive as we shall immediately see. What an organism feeds upon is negative entropy. Or, to put it less paradoxically, the essential thing in metabolism is that the organism succeeds in freeing itself from all the entropy it cannot help producing while alive.
WHAT IS ENTROPY?
What is entropy? Let me first emphasize that it is not a hazy concept or idea, but a measurable physical quantity just like the length of a rod, the temperature at any point of a body, the heat of fusion of a given crystal or the specific heat of any given substance. At the absolute zero point of temperature (roughly- 273Â°C) the entropy of any substance is zero. When you bring the substance into any other state by slow, reversible little steps (even if thereby the substance changes its physical or chemical nature or splits up into two or more parts of different physical or chemical nature) the entropy increases by an amount which is computed by dividing every little portion of heat you had to supply in that procedure by the absolute temperature at which it was supplied - and by summing up all these small contributions. To give an example, when you melt a solid, its entropy increases by the amount of the heat of fusion divided by the temperature at the melting-point. You see from this, that the unit in which entropy is measured is cal./Â°C (just as the calorie is the unit of heat or the centimetre the unit of length).
THE STATISTICAL MEANING OF ENTROPY
I have mentioned this technical definition simply in order to remove entropy from the atmosphere of hazy mystery that frequently veils it. Much more important for us here is the bearing on the statistical concept of order and disorder, a connection that was revealed by the investigations of Boltzmann and Gibbs in statistical physics. This too is an exact quantitative connection, and is expressed by
entropy = k log D,
where k is the so-called Boltzmann constant ( = 3'2983.10-24 cal./Â°C), and D a quantitative measure of the atomistic disorder of the body in question. To give an exact explanation of this quantity D in brief non-technical terms is well-nigh impossible. The disorder it indicates is partly that of heat motion, partly that which consists in different kinds of atoms or molecules being mixed at random, instead of being neatly separated, e.g. the sugar and water molecules in the example quoted above. Boltzmann's equation is well illustrated by that example. The gradual 'spreading out' of the sugar over all the water available increases the disorder D, and hence (since the logarithm of D increases with D) the entropy. It is also pretty clear that any supply of heat increases the turmoil of heat motion, that is to say, increases D and thus increases the entropy; it is particularly clear that this should be so when you melt a crystal, since you thereby destroy the neat and permanent arrangement of the atoms or molecules and turn the crystal lattice into a continually changing random distribution.
An isolated system or a system in a uniform environment (which for the present consideration we do best to include as a part of the system we contemplate) increases its entropy and more or less rapidly approaches the inert state of maximum entropy. We now recognize this fundamental law of physics to be just the natural tendency of things to approach the chaotic state (the same tendency that the books of a library or the piles of papers and manuscripts on a writing desk display) unless we obviate it. (The analogue of irregular heat motion, in this case, is our handling those objects now and again without troubling to put them back in their proper places.)
ORGANIZATION MAINTAINED BY EXTRACTING 'ORDER' FROM THE ENVIRONMENT
How would we express in terms of the statistical theory the marvellous faculty of a living organism, by which it delays the decay into thermodynamical equilibrium (death)? We said before: 'It feeds upon negative entropy', attracting, as it were, a stream of negative entropy upon itself, to compensate the entropy increase it produces by living and thus to maintain itself on a stationary and fairly low entropy level.
If D is a measure of disorder, its reciprocal, 1/D, can be regarded as a direct measure of order. Since the logarithm of 1/D is just minus the logarithm of D, we can write Boltzmann's equation thus:
- (entropy) = k log (1/D).
Hence the awkward expression 'negative entropy' can be replaced by a better one: entropy, taken with the negative sign, is itself a measure of order. Thus the device by which an organism maintains itself stationary at a fairly high level of orderliness ( = fairly low level of entropy) really consists in continually sucking orderliness from its environment. This conclusion is less paradoxical than it appears at first sight. Rather could it be blamed for triviality. Indeed, in the case of higher animals we know the kind of orderliness they feed upon well enough, viz. the extremely well-ordered state of matter in more or less complicated organic compounds, which serve them as foodstuffs. After utilizing it they return it in a very much degraded form - not entirely degraded, however, for plants can still make use of it. (These, of course, have their most powerful supply of ‘negative entropy’ in the sunlight.)
NOTE TO CHAPTER 6
The remarks on negative entropy have met with doubt and opposition from physicist colleagues. Let me say first, that if I had been catering for them alone I should have let the discussion turn on free energy instead. It is the more familiar notion in this context. But this highly technical term seemed linguistically too near to energy for making the average reader alive to the contrast between the two things. He is likely to take free as more or less an epitheton ornans without much relevance, while actually the concept is a rather intricate one, whose relation to Boltzmann's order-isorder principle is less easy to trace than for entropy and 'entropy taken with a negative sign', which by the way is not my invention. It happens to be precisely the thing on which Boltzmann's original argument turned.
But F. Simon has very pertinently pointed out to me that my simple thermodynamical considerations cannot account for our having to feed on matter 'in the extremely well ordered state of more or less complicated organic compounds' rather than on charcoal or diamond pulp. He is right. But to the lay reader I must explain that a piece of un-burnt coal or diamond, together with the amount of oxygen needed for its combustion, is also in an extremely well ordered state, as the physicist understands it. Witness to this: if you allow the reaction, the burning of the coal, to take place, a great amount of heat is produced. By giving it off to the surroundings, the system disposes of the very considerable entropy increase entailed by the reaction, and reaches a state in which it has, in point of fact, roughly the same entropy as before.
Yet we could not feed on the carbon dioxide that results from the reaction. And so Simon is quite right in pointing out to me, as he did, that actually the energy content of our food does matter; so my mocking at the menu cards that indicate it was out of place. Energy is needed to replace not only the mechanical energy of our bodily exertions, but also the heat we continually give off to the environment. And that we give off heat is not accidental, but essential. For this is precisely the manner in which we dispose of the surplus entropy we continually produce in our physical life process.
This seems to suggest that the higher temperature of the warm-blooded animal includes the advantage of enabling it to get rid of its entropy at a quicker rate, so that it can afford a more intense life process. I am not sure how much truth there is in this argument (for which I am responsible, not Simon). One may hold against it, that on the other hand many warm-blooders are protected against the rapid loss of heat by coats of fur or feathers. So the parallelism between body temperature and 'intensity of life', which I believe to exist, may have to be accounted for more directly by van't Hoff's law, mentioned on p. 65: the higher temperature itself speeds up the chemical reactions involved in living. (That it actually does, has been confirmed experimentally in species which take the temperature of the surroundings.)