Sei sulla pagina 1di 4

TIBS 20 - JANUARY 2 99,5 REFLECTIONS

the hi~to~y of science. (Fo~ further bio-


°.. ¢ . ?: graphical details, see Rds 3-5.)
Schrod nge s What/s L fe
a 50oyear reflection I do not intel~d here to review the
many commentaries on What is Lifp?
would, howeve{; like to address (and
hopefully give some redress to) the
Erwin Schr6dingefs What bs Life? BoRzmann. The untimely death of question of the originadity of his central
(Cambridge University Press, 1944) is Boltzmann dictated a course of study ideas - a subject that has attracted
one of the most talked about books in for Schrhdinger under the master's much attention and that, some wou~d
tv,r,.nt~,.~,,o,.,.,,t
" "~" . . . . . .,,,. .~, .~,.l,.l'~C,..
"~""" The book capable proteges. His closest friend argue, is the basis of the book's legacy.
came under intense scrutiny at the from those days in Vienna was Franz The maiorit3, of the book's content is, at
centenary of Schrhdinger's birth in 1987 ('Fr~inzer) FrimmeL a botany student face value, devoted to the molecular
(reviewed in Ref. 1) and was revisited with whom he had long late-night dis- aspects of genetic action, as the author
at a recent symposium (see Ref. 2). cussions about the meaning of life. regarded the chromosome as the funda-
Discussion has focused largely on its Together, they read Richard $emon's mental directing element in the gener-
impact on the early development ol our 1904 book, Die Mneme als erhaltendes ation of life's order. Accordingly, most
understanding of chromosomal struc- Frinzip (The Mneme as Conservative of the commentaries on the book have
ture and of the concept of the 'genetic Principle), which left a lasting im- concentrated on the genetic aspects o:
code'. Reflecting on the course of the pression on the young SchrhdingeL its historical status. Max Perutz 6, foc
scientific unity of physics and biology Semon was a pupil of Ernst Haeckel and example, has professed cogently that
over the 50 years since its publication, the atomistic 'plastidular' view of living What is Life? contains a considerable
one might ask, 'Does the fixation on the systems. Semon's Die Mneme was de- amount of textbook information on
historical role of What is Life? in the rived from a psycho-lamarckian analogy hereditary mechanisms and mutations,
birth of molecular genetics represent a between memory and inheritance. It is that the book's 'backbone' is simply
myopic view of the author's philosophi- probably more than coincidence tha~ a paraphrase of a 1935 article
cal intent and shortchange the book's Lamarck's Philosophic Zoologique was by Timof~eff-Ressovsky, Zimmer and
biophysical legacy? Were the book's one of the few books Schrhdmger Delbriick7 (.entitled 'The Nature of
ideas original? Was Schrfdinger merely retained from his father's library (the Genetic Mutations and the Structure of
applying physics as a tool to analyse a remainder having been sold owing to the Gene'), and that Schr6dinger's lack
biological object; or, rather, was he pon- financial necessity). of knowledge of the relevant literature
dering an expanded natural philosophy From the writings, lectures and per- invalidates the originality of his views.
that would provide a physicalist basis sonal contacts of his later life as a The legitimacy of such claims aside,
fi)r biology?' The fiftieth anniversary of world-famous physicist, it is evident many commentators (reviewed in Re[s
the publication o[ What is Life? serves that Schrodinger maintained a sideline 1, 3-5, 8) have argued that the book,
as a fitting opportunity to (re)examine interest in biology. Much of his thinking with its elegantly written, although
the significance of the biophysical comes to a head in What ~ Life?, which speculative, popularist style, was influ-
issues raised by Schrhdinge~: was based ~)n a series of public lectures ential in attracting the early attention of
at Trinity College in Dublin. The text is physicists and chemists, as well as biol-
A biogcaphlcalprelude an amalgam of the statistical physics ogists, to the paramount biological
Schr6dinger was certainly not the and thermodynamics of the author's problem of the molecular basis of gene
only theoretician from the golden turn-of-the-century training in Vienna, action. James Watson 9, for example,
period of physics in the early twentieth the quantum physics of his latter-day recalled '...I came back to the
century to delve into the science of profession, and his early conviction University of Chicago and spotted the
biology. Niels Bohr, whose father was towards biological order, evolution and tiny book What is Life? by the theoreti-
a physiologist, is a notable example; hea'edity as deterministic Darwinian cal physicist Erwin Schr6dinger. In that
but of the pioneers from that ger- processes. little gem, Schr6dinger said that the
minal realm of relativity and quan- The Dublin lectures were very popu- essence of life was the gene. Up until
tum physics, I would suggest that lar. However, the book elicited con- then I was interested in birds. But then
Schrfdinger had perhaps the deepest troversy even before its publication. thought, well, if the gene is the essence
interest in the physicalist ramifications In preparing the written version, of life, ! want to know more about it.
of the life sciences. His appreciation of Schr6dinger added an epilogue 'On And that was fateful because, other-
biology can be traced to frequent child- Determinism and Free Will', which con- wise, ! would have spent my life study-
hood discussions with his father, whose tained a denunciation of Western phil- ing birds, and no one would have heard
avocation was botany. (What is Life? osophy/religion and an espousal of of me.'
is dedicated to the memory of his Vedantic beliefs. Objection to this Moving away from the oft-discussed
parents.) Schrfdinger read Darwin's attack on the Church resulted in a col- role of What is Life? in the history of
The Origin of Species at an early age and lapse in the original plan for Cahill & genetics, I should like to consider in
became enthralled by the mechanistic Co. in Dublin to publish the book. some depth Schr6dingefs application of
view of the living world. Through a friend, the physical chemist statistical physics and thermodynamics
Schrfdinger entered the University Frederick Donnan, Schr6dinger got the to biology. His position was that 'it is in
of Vienna in 1906, hoping to pursue text published by Cambridge University relation to the statistical point of view
physics under the great Ludwig Press; and thus did What is Life? enter that the structure of the vital parts of

© 1995,ElsevierScienceLtd 0968.--0004/95/$09.50 45
S "rIBS 20 - JANUARY .~995

SchrSdlnger's concentration on the that 'it is dangerous to apply thermo-


rationality of ordered cellular process- dynamic arguments directly' to cellular
es from the standpoint of statistical processes. Instead, Peters suggested
physics was incisive, but (at the risk of that 'owing to the micro-heterogeneous
imparting a whiggish historiography*) nature of the system, surface effects
we now see his analysis as more appro- take precedence over ordinary statisti-
priate to the molecular characteriz- cal, mass action relationships, and
ation of the phenotype rather than to become in the ultimate limR respon-
that of the genotype. In Schr6dinger's sible for the integration of the whole
day, little was known empirically about and therefore the direction of activi-
the structural organization of cellular ties.' Crossing paths with Schr6dinger,
metabolism. Also, we should be Peters, taking 'the "gene" theory as a
reminded that biologists (especially in guide,' speculated that, 'if we isolate in
the early twentieth century and, to thought the instant...at which the
some extent, et,en today) tend to "gene" first commences to exert its
regard subcellu|ar processes according directive effect .... ' we should find that
to simplistic nineteenth-century physi- the protein-based cellular order begins
cochemical notions based on dilute, with the 'protamine' portion ()f the
weak-electrolyte aqueous solutions and chromatin.
Rgure 1 on gas-phase kinetics. Even so, early in We now know that larger eukaryotic
A portrait of Schr6dinger,taken by Lotte the twentieth century a number of theor- ceils are, indeed, laced with a dense
Meitner-Graf(c. :I.935). The Professor is
shown smoking his favorite Peterson eticians from such areas as biochem- array of fibrous proteinaceous elements,
pipe from Dublin. Reproduced from istry, cell biology and developmental as well as with spacious lipoprotein
Ref. 3 with permission from Cambridge biology were already contemplating membrane surfaces; and a multitude of
UniversityPress. seriously the ideas of spatial order and metabolic pathways has been found to
microenvironmentai deviation in the be associated with this cytomatdx ~3.
cellular machinery (reviewed in Refs The theoretical issue of m~croenviron-
living organisms differs so entirely from 11-13). In particular, the crystal mental statistical variation in metabolic
that of any piece of matter that we metaphor was being used at this time phenomena has continued to draw
physicists and chemists have ever as a 'field' concept by developmental interest even in later years (see, for
handled...'. Schr6dlnger illustrated his biologists (see Ref. 11). example, Ref. 15). Perh~,ps the most
biological concern by reference to the An interesting parallel to succinct way of characterizing local
so-called '~/n rule', whereby physlco- SchrSdinger's quest for understanding biochemical flow processes in living
chemical laws manifest an inaccuracy of the physical basis of cellular order, cells is to refer to them in a modified
with a relative error of the order of but on the phenotyplc rather than the 'probabllistic' manner, as a diffusion
l/~ln, where n Is the number of mol. genotyplc side, may be seen in an early under a bias, where the bias is rendered
ecules resl~aslble for the process. As paper entitled 'Surface Structure tn the by the ambient structural setting (see
most physical laws deal with systems Integration of Cell Activity', published Ref. 13). Such a specific designation
entailing very large numbers of par- in 1930 by the Oxford biochemist, was, In fact, suggested in 1925 by the
ticles, the error in predictability is Rudolph Peters =4.Akin to Schrfdlnger's mathematical biologist Alfred Lotka ~.
usually insignificant. Whence, he won. motivation, Peters felt that 'the prob- Schr~dinger also examined the gl,~bal
dered how the gene (or chromosome) - a lem of greatest significance Is that pre- problem of accounting for the or~,. "z-
single macromolecule - could deter- sented by the orderly, continuous and ation of the living state within the con-
mine dynamically 'the very orderly and coordinated direction of chemical pro- text of the second law of thermodynam-
lawful events within a living organism'. cesses' in the living ceil. With only ics. He maintained that such internal
Sehr6dinger's resolution to the para- limited cytological evidence available at order must be 'paid for' at the expense
dox led him to propose that life's the time, he argued for a =nechanism of degradation of substrates - 'negative
spatiotemporal order must be due, ulti- of 'directed chemical change' executed entropy' (or 'negentropy') - from the
mately, to the Inherent structural by enzyme populations associated with surroundings. In the immediate after-
properties of the chromosome, which a reflexive proteinaceous network ex- math of the publication of What is Life?,
he conceptualized as a heat-stable tending throughout the cell. Based on Sehr6dinger was criticized for not being
'aperiodic crystal', involving a 'minia- simple numerical calculations for the explicit with the technical details of the
ture code' for a one-to-one correspon. putative microcompartments posed by nature of open systems and with the
dence with the organismic plan. Thus, such ultrastructure, Peters pictured significance of Gibbs free energy in re-
he said, 'we are here obviously faced how fluctuations about some measured lation to negentropy. He retorted that
with events whose regular and lawful average value of cellular concentration the book was directed to a lay audience
unfolding is guided by a "mechanism" can be of great kinetic consequence for and that he was deliberately short on
entirely different from the =probability the localized biochemical events, in a detail, although in a later edition he
mechanism" of physics'. Schr6dinger's similar vein to Schr6dinger, he asserted provided an addendum on this point.
longstanding romance with biology While Schr6dinger lacked knowledge of
seems to reach a crescendo here, as he *As phrased by Ernst Mayrt°, 'whiggish' is the term some of the relevant biology, we can
mused over such a 'fantastic descrip- applied to that kind of biased, hindsighted history rest assured that, from his academic
tion, perhaps less becoming a scientist of science, 'in which every scientist is judged by the
extent of his [s/c] contribution toward the establish- upbringing under the shadow of
than a poet'. ment of our current interpretation of science.' Boltzmann in Vienna, he was well aware
T~BS 20 - JANUARY 1 9 9 5 REFLECTIONS
of the overall thermodynamic nature oi iee] the exciting hotism i~ presentoday maierial substance to ,+~imp~ephysical
the matter. physics and the urgency in its unifi- objects; ratheL ti~: t~eoe+eticai characo
Schr6dinger's thermodynamic reason- cation with biology. ter of space-time ~sociations for pro-
ing here was perspicacious. Although These hoiistic times stimulate many cesses to a physical foemat is reduced.
the basic principles had begun to thinkers to cross the barriers separat- Schr6dinger was, of course, caught in
appear in physics/chemistry textbooks ing the individual intellectual disci- the beginning of the ~mo!ecc~!arbiology'
(see, for example, Ref. 17) at the time, plines and to engage in liberai analogi- era, whictt comme~:.e(~ '-' , ih e a r n e s t
the subiect ol nonequilibrium thermo- cal thinking between them. 5chrSdinger shortly after the tu~r~,~-&e-century
dynamics was not well known, especially was a perceptive and inquisitive stu- rediscovery of MendeFa wm-k. Yet his
in the science of biology. Today, the dent of science during a similar period concern was not wit~ the nah~re of
thermodynamics of nonequilibrium pro- some 100 years ago. ! agree with the genes per se; rather, he w~i:ed to L~ow
cesses has generated a well-defined biographical speculation of W~lter the causal, dynamical basis of 'events'
scientific edifice that deals implicitly Moorea, that What is Life? had its (such as structures, organization arm
with the idea of negentropy as free- origins in the author's student days in hereditary transitions) in the ii~'ing
energy exchange across the boundary Vienna, in those late-night biological state. SchrSdinger's famous 'aper,:oaic
of open systems and internal dissi- discussions with Frlinzel FrimmeL As crystal' metaphor for chromosomal
pation (see, for example, Reis 18, 19). the nineteenth century turned into the structure should be interpreteO in this
Perhaps Schr6dinger's view came from twentieth century, there was an abun- dynamical sense. To a theoretical physi-
Boltzmann who, in the late nineteenth dance of books hailing the unity of sci- cist, the idea of a 'crystal' immediately
century, had lectured and written of liv- ence (reviewed in Ref. 13), among them conjures up mathematical symmetry
ing organisms as evading the second some by biologists such as Ernst groups. A hallmark of relativity and
law of thermodynamics by feeding on Haeckel, of whom we know SchrOdinger quantum physics in this century has
negative entropy (see Ref. 20). The was aware (see Ref. 3). Moreover, the been the elucidation of symmetry/
bioenergetic significance of BoRzmann's physicist Boltzmann, Schr6dinger's sci- asymmetry as the dynamical principle
depiction had been identified (and duly entific idol from his university period, underlying all the forces of Nature (see
cited), long before What is Life?, by was a Darwin enthusiast and had given Ref. 24). Pierre Curie 25, in the late 1800s,
Lotka (see Ref. 13). In 1940, Ludwig yon considerable notice to biological issues used the concepts of crystal symmetry
Bertalanffy (respected by many as the in the late nineteenth century (see groups in a dynamical manner (prob-
father of 'general systems theory'), Ref. 20). ably unknown to Schr6dingeO that,
independently had characterized living Moore a suggests that Schr6dinger's today, provides the theoretical basis for
systems in virtually the same terms as incentive for interdisciplinary transcen- our understanding of many kinds of
Schr6dinger (see Ref. 21). dence came from the Naturphilosophie coupled thermodynamic flow process
The central ideas in What is Life?, spirit of Goethe, in whom he was well in living cells (see Refs 13, 18). While we
when considered individually, would versed. The polymathic credo in cannot thank Schr6dinger solely for the
not seem all that original. Nevertheless, Goethe's On Generql Thc:ory tells all: 'If appearance of this paradigm in biology,
Schr6dinger's unique synthesis and you desire to reach out into the infinite, the prescience of his analogical thinking
physicalist insight gave notice, to a move in all directions in the finite.' is historically noteworthy.
wide and diverse scientific audience, of Goethe and Lamarck (whom, as noted • In looking at the plethora of works on
a number of important, but relatively above, Schr6dinger also read) were two the theoretical foundation of the unifi-
obscure, biophysical (as well as genetic) of the greatest universalists in the mod- cation of biology and physics being
issues. This role, in itself, is sufficient to ern history of science (see Ref. 22). written today, SchrSdinger would surely
ensure a venerable position for the What is Life? is not really about smile contentedly. Not only should he
book in the history of science. genes, nor is it about entropy. be revered for his fundamental role in
Schr6dinger, in [act, declares his pur- the physics revolution of the twentieth
Fr0m ideas i0 ih0ughts pose in the opening pages. After empha- century, but he should be remembered
The true legacy of What is Life? is to sizing 'the keen longing for unified, all- in the history of science as one of the
be realized by reflecting not so much on embracing knowledge' and apologizing first pioneers from the early stage of
the author's 'ideas' but on the back- to the reader for his 'second-hand and that revolution to extend the physical-
ground 'thoughts' that may have driven incomplete knowledge' of biology in the ist implications ('albeit,' as he said,
him to associate the ideas. At present, Preface, he states on the first page of • 'with secondLand and incomplete
there is a communal feeling of unity the first chapter his focus, 'at the risk of knowledge') to other areas.
(perhaps a fin-de.stale euphoria?) making fools of ourselves,' on the fun- Despite the phenomenal successes
amongst many scientists, especially damental question: 'How can the events of theoretical physics and of molecular
those of a theoretical bent. As the twen- in space and time which take place biology, we are, alas, no closer to
tieth century (and an old millennium) within the spatial boundary of a living answering the ultimate question What is
comes to a close, bookshops are replete organism be accounted for by physics Life? than in Schr6dinger's day. With
with titles beginning with The End of..., and chemistry?' (Schr6dinger's italics). time, the explanation becomes more
from all fields of human endeavor. Many The true significance of What is Life? is elusive and metaphysical. Dare we infer
prominent physicists are now boldly to be understood within the realm of that the author sensed the uncertainty
asserting that a 'theory of everything' physicalism, a philosophical program, of himself, in adding the controversial epi-
unification that accords a special rela- logue? As voiced by Lotka16, a contem-
(the so-called 'TOE') is at hand. One has
only to open the pages, for example, of tional privilege to physics (see Ref. 23). porary of Schr6dinger, '(in) searching
Paul Davies' book The New Physics Within this paradigm, the natural world for the essential characteristics of life,
is not analysed by merely reducing those that should finally and conclusively
(Cambridge University Press, 1989), to
47
: i _ ~ _

BOOKREVIEW TIBS 2 0 - JANUARY 1 9 9 5

distinguish ~he living from the non- 8 Symonds, N. (1986) O. Rev. Biol. 61, 19 Westerhoff, H. V. and Van Dam, K. (1987)
221-226 Thermodynamicsand Control of Biological Free-
living, are we not just searching for the 9 Watson, J. (1993) Science261, Energy Transduction, Elsevier
thing in nature that should correspond 1812-1813 20 Broda, E. (1983) LudwigBoltzmann:
to a word in our vocabulary? Are we not 10 Mayr, E. (1982) The Growth of BS.ological Man-Physicist-Philosopher,
hunting the Jabberwock?' Thought, Harvard University Press Ox BOWPress
11 Haraway,D. J. (1976) Crystals, Fabrics, and 21 Davidson, M. (1983) UncommonSense:
ReMs: Metaphors of Organicismin Twentieth- The Life and Thoughtof Ludwig yon Bertalanffy,
A©know~lgemnt Century DevelopmentalBiology,Yale University ]'archer
I am grateful to D. B. Kell and J. A. Press 22 Welch, G. R. 'Goethe's Gestalt, Bi/dung, and
12 Witkowski, J. A. (1987) Medical History 31. Urph&nomenin Biology: A Twentieth-Century
Witkows~J for helpful comments.
247-268 Physicalist View,' in GoetheScienziato(Grieco,
13 Welch, G. R. (1992) Prog. Biophys. Mot. Biol. A., ed.), Einaudi (in press)
References 57, 71-128 23 Poland, J. (1994) Physicalism:
i Welch. G. R. (1989) BioEssays1L 187-190 14 Peters, R. A. (1930) Trans. FaradaySoc. 26, The PhilosophicalFoundations,
2 Gnaiger, E. et aL (1994) What is Controlling 797--822 Oxford University Press
Life?, ;nnsbruck University Press ,t5 Smeach, S. C. and Gold, H. J. (1975) J. Theor. 24 Pageis, H. R. (1985) Perfect Symmetry,
3 Moore, W, (1989) SchrSdinger:Life and Biol. 51, 59-96 Simon & Schuster
Thought, Cambridge University Press 16 Lotka, A. J. (1925) Elementsof Physical Biology, 25 Curie, P. (1908) Oeuvresde Pierre Curie,
4 Yoxen, E. J. (1979) Hist. Sci. 17, 17-52 Williams & Wilkins Gauthier-Villars
5 Witkowski, J. A. (1986) Trends8iochem. Sci. 17 De Donder, T. and Van Rysselberghe, R (1936)
11, 266-268 ThermodynamicTheoryof Affinity, Stanford
6 Perutz, M. F. (1987) Nature326, 555-558
G. RICKEY WELCH
University Press
7 TimoE.=eff-Ressovsky,N. W., Zimmer, K. G. and 18 Caplan, S. R. and Essig, A. (1983)
Delbn3ck, M. {1935) Nachrichten aus tier Bioenergetics and Linear Nonequilibrium Department of Biological Sciences, University
B,ologie der Gese#schaflder Wissenschaften Thermodynamics,Harvard University of New Orleans, New Orleans, LA 70148,
G6ttingen 1, 189-245 Press USA.
.... L

The teenage weakling and has an appropriate number of clear


illustrations to make the points that the
because, if the book were much, much
longer, it wo_,dd look like a traditional text
other stories authors have in mind. Other strengths
include its breadth, clinical relevance,
and lose its appeal. It is a distinct
advantage to be able to put the book
and the mostly up-to-date, clear down for a period of time and pick it up
Biochemistry for the Medical explanations. Students in the health later to read any of the cases that one
sciences should certainly lind it highly may find interesting at that time.
Sciences: An Integrated Case instructive and interesting. I do not think There are, however, some points I
Approach students or faculties should be, or will be, would have wanted stressed in a book
'put off' by the selection of cases, some of that should 'make students think'. For
edited by S. J. Higgins, A. J. T~tmer and which describe rare diseases. example, quantitation is underdeveloped.
E. J. Wood, Longman, i 9 9 4 . £ 1 4 , 9 9 Before discussing my interpretation of in case one, the teenage weakling, it is
(xiii + 20:t pages) ISBN 0 5 8 2 1 0 1 2 9 8 the weaknesses, a brief comment about necessary to know that tlux through
the layout. The cases are presented pho~vholructokinase-1 has to be 18-fold
This is a self-study book that should succinctly. An introduction to the higher to regenerate the same amount of
attract the interest of students in the pi :)blem is provided in usually less than ATP anaerobically versus aerobically. If
health sciences, it might help them one page. This is followed by a concise the student appreciated this, it would
develop problem-solving skills and outline of the problem, including all the help explain why the patient doesn't have
curiosity, and allow them to avoid the relevant data. Questions are then asked, symptoms at rest or during aerobic
traditional barriers between basic science some being more usefu! than others, but exercise. Moreover, in areas where I have
disciplines and between basic versus most are pertinent. The authors choose more detailed knowledge, i was less
clinical departments. (wisely in my opinion) not to p=ovide happy about some of the discussion (for
it is now quite clear that the emphasis specific answers, but provide a example, case ten, acid excretors), but
in medical education has charged. Gone commentary that addresses all the key this again is not a major criticism.
axe the 'day-long' lectures and the passive elements with respect to the questions. As one who has a great interest in
style of education. The responsibility for To stimulate the student, further teaching basic sciences to medical
education is now translerred, to a large (unanswered) questions are provided. students, and one who has attempted to
degree, from the faculty to the student, Many of these require detailed achieve a somewhat similar aim l, overall 1
Further, there is a great need for information, but are, nevertheless, am very enthusiastic about this book. !
introducing clinical relevance and important questions, in the final section, would recommend it highly to teaching
motivation for students to seek the authors attempt to encourage faculties to help ~elect and develop case
information, in this light, this text seems integration, both horizontally between material, i v,ould make students aware of
very timely. The medical school faculty the basic sciences and vertically to more this book and encourage them to read it.
must search for or create new ways to clinical areas; this is the only area that is
present the basic sciences, ways that not sufficiently developed to the high Reference
capture the potential enthusiasm of the level of the remainder of the book. Finally, 1 Halperin, M. L. and Rolleston, F. S. (1993)
student while avoiding over-burdening excellent introductory references are Clinical Detective Stories: A Problem.Based
the student initially. Concepts and provided. Approach to Clinical Cases in Energyand
thinking patterns, rather than fact The strengths of this book are also its Acid-Base Metabolism, Portland Press
overload, are on everyone's minds. weaknesses. It covers so wide an area
Nevertheless, there are few texts available that each case is very 'thin'; it does not MITCHELL L. HALPERIN
to achieve these aims. cover any subject in depth, and reads
Biochemistry for the Medical Sciences is like a collection of interesting, but is the co-author of Clinical Detective Stories and is
a large step in the right direction. It is well relatively unrelated, case synopses. I do at St Michael's Hospital, 38 5huter Street, Toronto,
written, clearly presented, concise and not find this to be a major detraction Ontario, Canada M5B 1A6.
48

Potrebbero piacerti anche