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PHYSICS IN MOLECULAR BIOLOGY
Cambridge University Press has no responsibility for the persistence or accuracy of s
for external or third-party internet websites referred to in this publication, and does not
guarantee that any content on such websites is, or will remain, accurate or appropriate.
Contents
v
Preface
vii
viii Preface
Kim Sneppen is particularly grateful for the hospitality of KITP at the University
of California, Santa Barbara, where material for part of this book was collected
during long visits at programs on Physics in Biological Systems in winter/spring
2001 and 2003.
I, Kim, thank my infinitely wise and beautiful wife Simone for patience and love
throughout this work, and my children Ida, Thor, Eva and Albert for putting life
into the right perspective.
Introduction
This book covers some subjects that we find inspiring when teaching physics stu-
dents about biology. The book presents a selection of topics centered around the
physics/biology/chemistry of genes. The focus is on topics that have inspired math-
ematical modeling approaches. The presentation is rather condensed, and demands
some familiarity with statistical physics from the reader. However, we attempted
to make the book complete in the sense that it explains all presented models and
equations in sufficient detail to be self-contained. We imagine it as a textbook for
the third or fourth years of a physics undergraduate course.
Throughout the book, in particular in the introductions to the chapters, we have
expressed basic biology ideas in a very simplified form. These statements are meant
for the physics student who is approaching the biological subject for the first time.
Biology textbooks are necessarily more descriptive than physics books. Our sim-
plified statements are meant to reduce this difference in style between the two
disciplines. As a consequence, the expert may well find some statements objection-
able from the point of view of accuracy and completeness. We hope, however, that
none is misleading. One should think of these parts as first-order approximations
to the more complicated and complete descriptions that molecular biology text-
books offer. On the other hand, the physical reasoning that follows the simplified
presentation of the biological system is detailed and complete.
The book is not comprehensive. Large and important areas of biological physics
are not discussed at all. In particular we have not ventured into membrane physics
and transport across membranes, signal transmission along neurons and sensory
perception, to mention a few examples. While there are already excellent books
and reviews on all these subjects, the reason for our limited choice of topics is
more ambitious. The basic physics ideas that are relevant for molecular biol-
ogy can be learned on a few specific examples of biological systems. The ex-
amples were chosen because we find them particularly suited to illustrate the
physics.
1
2 Introduction
We have chosen to place the focus on genes, DNA, RNA and proteins, and in
particular how these build a functional system in the form of the λ-phage switch.
We further elaborate with some larger-scale examples of molecular networks and
with a short overview of current models of biological evolution. The overall plan of
the book is to proceed from simple systems toward more complex ones, and from
small-scale to large-scale dynamics of biological systems.
Chapter 1 gives some impression of important ideas in biology. To be more
precise, the chapter summarizes those concepts which, we think, strike a physicist
who approaches the field, either because they have no counterpart in physics, or, on
the contrary, because they are all too familiar. The chapter grew out of discussions
with biologists, and we normally use it as a first introductory lecture when we
give the course. Of the subsequent chapters, we regard Chapter 7 on the λ-phage in
E. coli as especially central: it deals with the interplay between elements introduced
earlier in the book, and it contains a lot of the physics reasoning that the book is
meant to teach.
In Chapter 2 we describe the physics of polymer conformations, emphasizing the
interplay between energy and entropy and examining both the behavior of extended
polymers and how compact configurations may be reached. In the next chapters
we introduce and discuss the most important biological polymers: DNA, RNA and
proteins. Although the covalent bonds forming the polymer backbone have binding
energies G > 1 eV, the form and function of these biomolecules is associated to
the much weaker forces perpendicular to the polymer backbone. These interactions
are of order kB T , and it is the combined effect of many of these forces that forms
the functional biomolecule. In Chapters 3–5 we characterize the stability of DNA,
RNA and proteins, with emphasis on the cooperativity responsible for this stability.
Biological molecules can be used for various types of computations. Chapter 3
includes a section on DNA computation and DNA manipulation in the laboratory.
This is in part a continuation of Chapter 2 (reptation), and also an introduction to
the computational aspects of molecular replication (the PCR reaction). Chapters
4–6, on the other hand, focus on proteins and protein folding and thus the functional
aspects are left to subsequent chapters. In this book we have addressed in consider-
able detail one of these aspects, namely how a protein may control the production
of another protein (Chapter 7). As we explain in Chapter 7, genetic control in-
volves mechanisms associated to both equilibrium statistical mechanics and to the
timescales involved in complex formation and disruption. Topics in this chapter
include a discussion of cooperativity, of target location by diffusion, of timescales
in a cell and of stability of expressed genetic states.
Chapter 7 also forms a microscopic foundation for the large-scale properties of
molecular networks, which we discuss in Chapter 8. Chapter 8 thus continues the
subject of genetic regulation and molecular networks, in part by venturing into the
Introduction 3
heat shock mechanism. This shows that protein folding is also a control mechanism
in a living cell, and it introduces a type of genetic regulation that was not treated in
the previous chapter: σ sub-units of RNAp, which control the expression of larger
classes of genes. Chapter 8 also discusses the larger-scale properties of genetic
regulatory networks, introducing a few recent physics attempts at modeling these.
Chapter 9 discusses evolution, with emphasis on the interplay between random-
ness and selection from the smallest to the largest scales. The chapter introduces
concepts such as neutral evolution, hill climbers and co-evolution, and uses these
concepts to discuss questions related in part to the concept of punctuated equilib-
rium, and in part to the origin of life in the form of autocatalytic networks. Thus
Chapter 9 introduces some simple models that allow us to discuss the nature of
the history leading to the emergence of life, and in particular aims at stressing
the importance of interactions and stochastic events on all scales of the biological
hierarchy.
In the Appendix we have a short introduction to statistical mechanics, including
the fluctuation–dissipation theorem and the Kramers escape problem; it is meant
to render the book self-contained from the point of view of the physics.
1
What is special about living matter?
Kim Sneppen & Giovanni Zocchi
Life is self-reproducing, persistent (we are ∼ 4 × 109 years old), complex (of the
order of 1000 different molecules make up even the simplest cell), “more” than the
sum of its parts (arbitrarily dividing an organism kills it), it harvests energy and
it evolves. Essential processes for life take place from the scale of a single water
molecule to balancing the atmosphere of the planet. In this book we will discuss
the modeling and physics associated, in particular, to the molecules of life and how
together they form something that can work as a living cell. First we briefly review
some basic concepts of living systems, with emphasis on what makes biological
systems so different from the systems that one normally studies in physics.
Conceptually, molecular biology has provided us with a few fundamen-
tal/universal mechanisms that apply over and over. Some concepts, like evolution,
do not have counterparts in physics. Others, like the role of stochastic processes,
are, on the contrary, quite familiar to a physicist.
(1) Biology is the result of a historical process. This means that it is not possible to
“explain” a biological system by applying a few fundamental laws in the same way
that is done in physics. A hydrogen atom could not be different from what it is, based
on what we know of the laws of nature, but an E. coli cell could. In evolution, it is much
easier to modify existing mechanisms than to invent new ones. Thus on evolutionary
timescales nearly everything comes about by cut and paste of modules that are already
working. We will end the book with a chapter dedicated to evolutionary concepts and
models.
(2) The molecules of life are polymers. At the molecular scale, life is made of polymers:
DNA, RNA and proteins. Even membranes are built of molecules with large aspect
ratios. Perhaps mechanics at the nano-scale can work only with polymers, molecules
that are kept together by strong forces along their backbone, while having the property
of forming specific structures by utilizing the much weaker forces perpendicular to the
backbone. In molecular biology we witness nano-mechanics at work with polymers.
We will discuss polymers in Chapter 2, and thereby introduce concepts necessary
4
What is special about living matter? 5
for understanding DNA (Chapter 3), proteins (Chapters 4–5) and polymers in action
(Chapter 6).
(3) Genetic code. Information is maintained on a one-dimensional, double-stranded DNA
molecule, which will be discussed in Chapter 3. Thus the one-dimensional nature of
the information mirrors the one-dimensional nature of the polymers that make life
work. The DNA strands open for copying and transcribing, by separating the double-
stranded DNA into two single strands of DNA that each carry the full information.
The copying is done by DNA polymerase using the complementarity of base pairs.
Similarly the genetic code is read by RNA polymerase and ribosomes that again use
the matching of complementary base pairs to translate codons into amino acids. This
is usually summarized in terms of the central dogma
This is highly simplified: proteins modify other proteins, and most importantly proteins
provide both positive and negative feedback on all the arrows in (1.1). If one has only
DNA in a test tube, nothing happens. One needs proteins to get DNA → RNA, etc.
Then Eq. (1.1) should be supplemented at least by an arrow from protein to DNA.
Thus it is not always clear where the start of this loop is, and the whole scheme has to
be extended to the complicated molecular networks discussed in Chapter 8.
(4) Computation. A living cell is an incredible information-processing machine: an
E. coli transcribes about 5 × 106 genes during 1/2 h, i.e. about 10 Gb/h of informa-
tion. All this within a 1 µm3 cell, coded by about 5 × 106 base pairs. The information
density far outnumbers that in any computer chip, and even a million E. coli occupy
A T
C G
T A
G C
T A
T A
A T
A T
C G
T A
G C
much less space than a modern CPU, thus beating PCs on computation speed as well.
The levels of computation in a living system increase when one goes to eukaryotes and
especially to multi-cellular organisms (where each cell must have encoded awareness
of its social context also). The simplest organisms (e.g. the prokaryote M. pneumono-
miae with 677 genes) can manage essentially without transcription control. Larger
genome size prokaryotes typically need a number of control units that grow with the
square of number of genes. We discuss modeling of processes within living cells in
Chapter 7 and, to some extent, also in Chapters 6 and 8.
(5a) Life is modular. It is build of parts that are build of parts, on a wide range of scales.
This facilitates robustness: if a process doesn’t work, there are alternative routes to
replace it. Molecular-scale examples include the secondary, tertiary and quaternary
structures of proteins (complexes of proteins); they may include network modules,
such as sub-cellular domains, that each facilitate an appropriate response to external
stimuli. Most importantly, the minimum independent living module is the cell.
(5b) Life is NOT modular. Life is more than the sum of its parts. Removing a single protein
species often leads to death for an organism. Another observation is that the number of
regulatory proteins for prokaryotes increases with the square of the number of proteins
that should be regulated. Thus regulatory networks are an integrated system, and not
modular in any simple way. This is the subject for the chapter on networks.
(6) Stochastic processes play an essential role from molecules to cells; in particular,
they include mechanisms driven by Brownian noise, trial-and-error strategies, and the
individuality of genetically identical cells owing to their finite number of molecules.
An example of a trial-and-error mechanism is microtubule growth, attachment and
collapse (see Chapter 6). Individuality of cells has been explored by individual cell
measurements of gene expression, and variability of cell fate has been associated
with fluctuations in gene expressions. An example of such stochasticity includes the
lysis–lysogeny decision in temperate phages; see Chapter 7.
(7) Biological physics is “kB T -physics”. The relevant energy scale for the molecular
interactions that control all biological mechanisms in the cell is kB T , where T is room
temperature and k is the Boltzmann constant (kB NA = R, where NA is Avogadro’s
number and R is the gas constant; 1 kB T = 4.14 × 10−14 ergs = 0.62 kcal/mole at
T = 300 K ). This is not true for most of the systems described in a typical physics
curriculum, for example:
r the hydrogen atom, with an energy scale ∼ 10 eV, whereas kB Troom 1/40 eV;
r binding energies of atoms in metals; covalent bonds: energy ∼ 1 eV;
r macroscopic objects (pendulum, billiard ball), where even a 1 mg object moving
with a speed of 1 cm/s has an energy ∼ 10−10 J ∼ 109 eV (1 eV = 1.602 × 10−19 J).
The approach is therefore different. For example, in the solid state one starts with a
given structure and calculates energy levels. Thermal energy may be relevant to kick
carriers in the conduction band, but kB T is not on the brink of destroying the ordered
structure.
Soft-matter systems often self-assemble in a variety of structures (e.g. amphiphilic
molecules in water form micelles, bilayers, vesicles, etc.; polypeptide chains fold to
Further reading 7
form globular proteins). These ordered structures exist in a fight against the disruptive
effect of thermal motion. The quantity that describes the disruptive effect of thermal
motion is the entropy S, a measure of microscopic disorder that we review in the
Appendix. So for these systems energy and entropy are both equally important, and
one generally considers a free energy F = E − T S. The language and formalism of
thermodynamics are effective tools in describing these systems. For example: free-
energy differences are just as “real” as energy differences; therefore entropic effects
can result in actual forces, as we discuss in Chapter 2.
Further reading
Berg, H. C. (1993). Random Walks in Biology. Princeton: Princeton University Press.
Boal, D. H. (2002). Mechanics of the Cell. Cambridge University Press.
Bray, D. (2001). Cell Movements: From Molecules to Motility. Garland Publishing.
Crick, F. H. C. (1962). The genetic code. Sci. Amer. 207, 66–74; Sci. Amer. 215, 55–62.
Eigen, M. (1992). Steps Towards Life. Oxford University Press.
Godsell, D. (1992). The Machinery of Life. Springer Verlag.
Gould, S. J. (1991). Wonderful Life, The Burgess Shale and the Nature of History. Penguin.
Howard, J. (2001). Mechanics of Motor Proteins and the Cytoskeleton. Sinauer Associates.
Kauffman, S. (1993). The Origins of Order. Oxford University Press.
Lovelock, J. (1990). The Ages of Gaia. Bantam Books/W. W. Norton and Company Inc.
Pollack, G. H. (2001). Cells, Gels and the Engines of Life. Ebner & Sons Publishers.
Ptashne, M. & Gann, A. (2001). Genes & Signals. Cold Spring Harbor Laboratory.
Raup, D. (1992). Extinction: Bad Genes or Bad Luck? Princeton University Press.
Schrödinger, E. (1944). What is Life? Cambridge University Press.
2
Polymer physics
Kim Sneppen & Giovanni Zocchi
Living cells consist of a wide variety of molecular machines that perform work
and localize this work to the proper place at the proper time. The basic design idea
of these nano-machines is based on a one-dimensional backbone, a polymer. That
is, these nano-machines are not made of cogwheels and other rigid assemblies of
covalently interlocked atoms, but rather are based on soft materials in the form
of polymers – i.e. one-dimensional strings. In fact most of the macromolecules
in life are polymers. Along a polymer there is strong covalent bonding, whereas
possible bonds perpendicular to the polymer backbone are much weaker. Thereby,
the covalent backbone serves as a scaffold for weaker specific bonds. This opens
up the possibility (1) to self-assemble into a specific functional three-dimensional
structure, (2) to allow the machine parts to interact while maintaining their identity,
and (3) to allow large deformations. All three properties are necessary ingredients
for parts of a machine on the nano-scale. In this chapter we review the general
properties of polymers, and thus hope to familiarize the reader with this basic
design idea of macromolecules.
Almost everything around us in our daily life is made of polymers. But despite
the variety, all the basic properties can be discussed in terms of a few ideas. Some
of these properties are astounding: consider a metal wire and a rubber band. The
metal wire can be stretched about 2% before it breaks; its elasticity comes from
small displacements of the atoms around a quadratic energy minimum. The rubber
band, on the other hand, can easily be stretched by a factor of 4. Clearly its elasticity
must be based on an entirely different effect (it is in fact based on entropy); see also
Fig. 2.1.
Polymers are long one-dimensional molecules that consist of the repetition of
one or a few units (the “monomers”) bound together with covalent bonds. You
can think of beads on a string. Figure 2.2 shows three examples; the first two are
synthetic polymers, the third represents the primary structure of proteins. What is
radically different between these molecules and all others is that the number of
8
Polymer physics 9
One-dimensional backbone
Three-dimensional structure
Cl Cl
R1 O R3 polypeptide
CH NH C CH
NH C CH NH C
O R2 O
monomers, N , is large, typically N ∼102 −104 (but note that for DNA N can be
∼108 ). The single most dramatic consequence is that the molecule becomes flexible.
We normally think of the relative motion of atoms within a small molecule, say
CO2 , in terms of vibrational modes. A polymer, however, can actually bend like a
string! There are more consequences. Perpendicular to the strong (covalent) forces
along the one-dimensional backbone, weaker forces may come into play; forces
that would be insignificant if the atoms were not brought together by the backbone
bonds. But given that the backbone forces these monomers together, the cooperative
10 Polymer physics
Polymerization
Polycondensation
O O
R1 C OH + OH R2 R1 C O R2
+ H2O
binding of many of these weaker forces, both within the same molecule and between
different molecules, allows the enormous number of specific interactions found in
the biological world.
In this chapter we will study the simplest polymers, consisting of many identical
monomers (“homopolymers”). This allows us to gain insight into the interplay
between the one-dimensional polymer backbone and the possible three-dimensional
conformations of the molecule.
Polymers are formed by polymerization (e.g. polyethylene) or by polyconden-
sation (e.g. polypeptides); see Fig. 2.3. The single most important characteristic
of polymers is that they are flexible. The simplest mechanism for their flexibil-
ity comes from rotations around single bonds. Figure 2.4 shows three links of,
say, a polyethylene chain; the C atoms are at the vertices and the segments depict
the C–C bonds. The bond angle θ is fixed, determined by the orbital structure of
the carbon, but φ rotations are allowed. As a result, on a scale much larger than the
monomer size a snapshot of the polymer chain may look as depicted on the right in
the figure, i.e. a coil with random conformation. For other polymers, for example
double-stranded DNA, the chemical structure does not allow bond rotations as in
Exploring the Variety of Random
Documents with Different Content
replied with princely indignation, that if honour were banished from
every other residence, it ought to find refuge in the breasts of kings.
Luther was permitted to retire from the diet; but he had not
proceeded far on his return when he was surprised by a number of
armed men, and carried away into captivity. It was an act of friendly
violence. A temporary concealment was thought necessary for his
present security, and he was hastily conveyed to the solitary Castle
of Wartenburg. In the mean time the assembly issued the
declaration known in history as the “Edict of Worms,” in which the
Reformer was denounced as an excommunicated schismatic and
heretic; and all his friends and adherents, all who protected or
conversed with him, were pursued by censures and penalties. The
cause of papacy obtained a momentary, perhaps only a seeming
triumph, for it was not followed by any substantial consequences;
and while the anathematized Reformer lay in safety in his secret
Patmos, as he used to call it, the Emperor withdrew to other parts of
Europe to prosecute schemes and interests which then seemed far
more important than the religious tenets of a German Monk.
While Luther was in retirement, his disciples at Wittemberg, under
the guidance of Carlostadt, a man of learning and piety, proceeded
to put into force some of the first principles of the Reformation. They
would have restrained by compulsion the superstition of private
masses, and torn away from the churches the proscribed images.
Luther disapproved of the violence of these measures; or it may also
be, as some impartial writers have insinuated, that he grudged to
any other than himself the glory of achieving them. Accordingly,
after an exile of ten months, he suddenly came forth from his place
of refuge, and appeared at Wittemberg. Had he then confined his
influence to the introduction of a more moderate policy among the
reformers, many plausible arguments might have been urged in his
favour. But he also appears, unhappily, to have been animated by a
personal animosity against Carlostadt, which was displayed both
then and afterwards in some acts not very far removed from
persecution.
The marriage of Luther, and his marriage to a nun, was the event of
his life which gave most triumph to his enemies, and perplexity to
his friends. It was in perfect conformity with his masculine and
daring mind, that having satisfied himself of the nullity of his
monastic vows, he should take the boldest method of displaying to
the world how utterly he rejected them. Others might have acted
differently, and abstained, either from conscientious scruples, or,
being satisfied in their own minds, from fear to give offence to their
weaker brethren; and it would be presumptuous to condemn either
course of action. It is proper to mention that this marriage did not
take place till the year 1525, after Luther had long formally rejected
many of the observances of the Roman Catholic Church; and that
the nun whom he espoused had quitted her convent, and renounced
her profession some time before.
The war of the peasants, and the fanaticism of Munster and his
followers, presently afterwards desolated Germany; and the papal
party did not lose that occasion to vilify the principles of the
reformers, and identify the revolt from a spiritual despotism with
general insurrection and massacre. It is therefore necessary here to
observe, that the false enthusiasm of Munster was perhaps first
detected and denounced by Luther; and that the pen of the latter
was incessantly employed in deprecating every act of civil
insubordination. He was the loudest in his condemnation of some
acts of spoliation by laymen, who appropriated the monastic
revenues; and at a subsequent period so far did he carry his
principles, so averse was he, not only from the use of offensive
violence, but even from the employment of force in the defence of
his cause, that on some later occasions he exhorted the Elector of
Saxony by no means to oppose the imperial edicts by arms, but
rather to consign the persons and principles of the reformers to the
protection of Providence. For he was inspired with a holy confidence
that Christ would not desert his faithful followers; but rather find
means to accomplish his work without the agitation of civil disorders,
or the intervention of the sword. That confidence evinced the perfect
earnestness of his professions, and his entire devotion to the truth of
his principles. It also proved that he had given himself up to the
cause in which he had engaged, and that he was elevated above the
consideration of personal safety. This was no effeminate enthusiasm,
no passionate aspiration after the glory of martyrdom! It was the
working of the Spirit of God upon an ardent nature, impressed with
the divine character of the mission with which it was intrusted, and
assured, against all obstacles, of final and perfect success.
As this is not a history of the Reformation, but only a sketch of the
life of an individual reformer, we shall at once proceed to an affair
strongly, though not very favourably, illustrating his character. The
subject of the Eucharist commanded, among the various doctrinal
differences, perhaps the greatest attention; and in this matter Luther
receded but a short space, and with unusual timidity, from the faith
in which he had been educated. He admitted the real corporeal
presence in the elements, and differed from the church only as to
the manner of that presence. He rejected the actual and perfect
change of substance, but supposed the flesh to subsist in, or with
the bread, as fire subsists in red-hot iron. Consequently, he
renounced the term transubstantiation, and substituted
consubstantiation in its place. In the mean time, Zuinglius, the
reformer of Zuric, had examined the same question with greater
independence, and had reached the bolder conclusion, that the
bread and wine are no more than external signs, intended to revive
our recollections and animate our piety. This opinion was adopted by
Carlostadt, Œcolampadius, and other fathers of the Reformation,
and followed by the Swiss Protestants, and generally by the free
cities of the Empire. Those who held it were called Sacramentarians.
The opinion of Luther prevailed in Saxony, and in the more northern
provinces of Germany.
The difference was important. It was felt to be so by the reformers
themselves; and the Lutheran party expressed that sentiment with
too little moderation. The Papists, or Papalins (Papalini), were alert
in perceiving the division, in exciting the dissension, and in inflaming
it, if possible, into absolute schism; and in this matter it must be
admitted, that Luther himself was too much disposed by his
intemperate vehemence to further their design. These discords were
becoming dangerous; and in 1529, Philip, Landgrave of Hesse, the
most ardent among the protectors of the Reformation, assembled
the leading doctors of either party to a public disputation at
Marpurg. The particulars of this conference are singularly interesting
to the theological reader; but it is here sufficient to mention, without
entering into the doctrinal merits of the controversy, that whatever
was imperious in assertion and overbearing in authority, and
unyielding and unsparing in polemical altercation, proceeded from
the mouth and party of Luther; that every approach to humility, and
self-distrust, and mutual toleration, and common friendship, came
from the side of Zuinglius and the Sacramentarians. And we are
bound to add, that the same uncompromising spirit, which precluded
Luther from all co-operation or fellowship with those whom he
thought in error (it was the predominant spirit of the church which
he had deserted) continued on future occasions to interrupt and
even endanger the work of his own hands. But that very spirit was
the vice of a character, which endured no moderation or concession
in any matter wherein Christian truth was concerned, but which too
hastily assumed its own infallibility in ascertaining that truth. Luther
would have excommunicated the Sacramentarians; and he did not
perceive how precisely his principle was the same with that of the
church which had excommunicated himself.
Luther was not present at the celebrated Diet of Augsburg, held
under the superintendence of Charles V. in 1530; but he was in
constant correspondence with Melancthon during that fearful period,
and in the reproofs which he cast on the temporizing, though
perhaps necessary, negotiations of the latter, he at least exhibited
his own uprightness and impetuosity. The ‘Confession’ of the
Protestants, there published, was constructed on the basis of
seventeen articles previously drawn up by Luther; and it was not
without his counsels that the faith, permanently adopted by the
church which bears his name, was finally digested and matured.
From that crisis the history of the Reformation took more of a
political, less of a religious character, and the name of Luther is
therefore less prominent than in the earlier proceedings. But he still
continued for sixteen years longer to exert his energies in the cause
which was peculiarly his own, and to influence by his advice and
authority the new ecclesiastical system.
He died in the year 1546, the same, as it singularly happened, in
which the Council of Trent assembled, for the self-reformation and
re-union of the Roman Catholic Church. But that attempt, even had
it been made with judgment and sincerity, was then too late. During
the twenty-nine years which composed the public life of Luther, the
principles of the Gospel, having fallen upon hearts already prepared
for their reception, were rooted beyond the possibility of extirpation;
and when the great Reformer closed his eyes upon the scene of his
earthly toils and glory, he might depart in the peaceful confidence
that the objects of his mission were virtually accomplished, and the
work of the Lord placed in security by the same heaven-directed
hand which had raised it from the dust.
RODNEY.
Engraved by E. Scriven.
LORD RODNEY.
London, Published by
Charles Knight, Pall Mall
East.
He was soon raised to a more important sphere of action, being
named Commander-in-Chief at Barbadoes and the Leeward Islands,
in the autumn of 1761. No naval achievement of remarkable
brilliance occurred during the short period of his holding this
command: but the capture of the valuable islands of Martinique, St.
Lucia, and Grenada, bears testimony to the efficiency of the fleet
under his orders, and the good understanding between the land and
sea forces employed in this service. He was recalled on the
conclusion of peace in 1763. Eight years elapsed before he was
again called into service; a period fruitful in marks of favour from the
crown, though barren of professional laurels. He was created a
Baronet soon after his return; he was raised by successive steps to
the rank of Vice-Admiral of the Red; and he was appointed Governor
of Greenwich Hospital. This office he was required to resign on being
again sent out to the West Indies as Commander-in-Chief at Jamaica
in 1771. This was a period of profound peace: but the duties of
peace are often more difficult, and require more moral courage for
their discharge, than those of war. It is one of Rodney’s best claims
to distinction, that he suffered none under his command, or within
the sphere of his influence, to neglect their duties with impunity:
and in the mode of carrying on naval affairs then practised in the
West Indies, he found much ground for immediate interference, as
well as for representation and remonstrance to his superiors at
home. He earnestly desired to obtain the government of Jamaica;
but on a vacancy occurring in 1773, another person was appointed;
and he was recalled, and struck his flag at Portsmouth, September
4, 1774.
The next four years of Sir George Rodney’s life were much harassed
by pecuniary embarrassment. The habits of a sailor’s life are
proverbially unsuited to strict economy: and moving, when at home,
in the most fashionable society of London, it is no wonder that his
expenses outran his professional gains. He was compelled to retire
to Paris, where he remained until the American war afforded a
prospect of his being called into active service again. In May, 1778,
he was promoted to the rank of Admiral of the White: but it was not
till the autumn of 1779 that he was gratified by being re-appointed
to the command on the Barbadoes station. He sailed from Plymouth
December 29, to enter on the final and crowning scene of his glory.
At this time Spain and France were at war with England. The
memorable siege of Gibraltar was in progress, and a Spanish fleet
blockaded the Straits. The British navy was reduced unwarrantably
low in point of disposable force; and was farther crippled by a spirit
of disunion and jealousy among its officers, arising partly perhaps
from the virulence of party politics, and partly from the misconduct
of the Admiralty, which threatened even worse consequences than
the mere want of physical force. By this spirit Sir George Rodney’s
fleet was deeply tainted, to his great mortification and the great
injury of the country. At first, however, every thing appeared to
prosper. The fleet consisted of twenty-two sail of the line, and eight
frigates. Before Rodney had been at sea ten days, he captured
seven Spanish vessels of war, with a large convoy of provisions and
stores; and on January 16, near Cape St. Vincent, afterwards made
memorable by a more important action, he encountered a Spanish
fleet commanded by Don Juan de Langara, of eleven ships of the
line and two frigates. The superiority of the British force rendered
victory certain. Five Spanish ships were taken, and two destroyed;
and had not the action been in the night, and in tempestuous
weather, probably every ship would have been captured. These at
least are the reasons which Rodney gave in his despatches, for not
having done more: in private letters he hints that he was ill-
supported by his captains. Trifling as this success would have
seemed in later times, it was then very acceptable to the country;
and the Admiral received the thanks of both Houses of Parliament.
The scandalous feeling of jealousy of their commander, ill-will to the
ministry, or whatever other modification of party spirit it was, which
could prevent brave men (and such they were) from performing
their duty to the utmost in the hour of battle, broke out again with
more violence when Rodney next came within sight of the enemy.
This was near Martinique, April 17, 1780, about a month after his
arrival in the West Indies. The French fleet, commanded by the
Comte de Guichen, was slightly superior in force. Rodney’s intention
was to attack the enemy’s rear in close order and with his whole
strength; but his captains disobeyed his orders, deranged his plan,
and careless of the signals for close action, repeatedly made, kept
for the most part at cautious distance from the enemy. His own ship,
the Sandwich, engaged for an hour and a half a seventy-four and
two eighty-gun ships, compelled them to bear away, and broke
completely through the enemy’s line. Not more than five or six ships
did their duty. Had all done it, the victory over De Grasse might have
been anticipated, and the end of the war accelerated perhaps by two
years. In his despatches Rodney censured the conduct of his
captains; but the Admiralty thought proper to suppress the passage.
In his private letters to Lady Rodney, he complains bitterly. One only
of his captains was brought to trial, and he was broken. That ampler
justice was not done on the delinquents, is to be explained by the
difficulty of finding officers to form courts martial, where almost all
were equally guilty. But this partial severity, with the vigorous
measures which the Admiral took to recall others to their duty,
produced due effect, and we hear no more of want of discipline, or
reluctance to engage. For this action Rodney received the thanks of
the House of Commons, with a pension for himself and his family of
£2000 per annum.
Nothing of importance occurred during the rest of the spring; and
De Guichen having returned to Europe, Rodney sailed to New York,
to co-operate, during the rainy season in the West Indies, with the
British forces engaged in the American war. In November he
returned to his station. In the course of the autumn he had been
chosen to represent Westminster without expense, and had received
the Order of the Bath. The commencement of the following year was
signalized by acts of more importance. The British ministry had been
induced to declare war against Holland; and they sent out immediate
instructions to Rodney, to attack the possessions of the states in the
West Indies. St. Eustatius was selected for the first blow, and it
surrendered without firing a shot. Small and barren, yet this island
was of great importance for the support which it had long afforded
to the French and Americans under colour of neutrality, and for the
vast wealth which was captured in it. In the course of the spring,
Demerara, Essequibo, and Berbice, with the French island of St.
Bartholomew, were also taken.
In the autumn, Rodney returned to Europe for the recovery of his
health. He was received with distinguished favour by the King, and
with enthusiasm by the people, and during his stay, was created
Vice-Admiral of Great Britain, in the place of Lord Hawke, deceased.
He returned in the middle of January, being invested with the
command of the whole West Indies, not merely the Barbadoes
station, as before. The situation of affairs at this time was very
critical. The French fleet, commanded by the Comte de Grasse,
consisted of thirty-three sail[6] of the line, two fifty-gun ships and
frigates, with a large body of troops, and a train of heavy cannon on
board. A powerful Spanish fleet was also in the West Indies. It was
intended to form a junction, and then with an overwhelming force of
near fifty sail of the line, to proceed to Jamaica, conquer that
important island, and one by one to reduce all the British colonies.
Joseph Louis Lagrange was born at Turin, January 25th, 1736. His
great-grandfather was a Frenchman, who entered into the service of
the then Duke of Savoy; and from this circumstance, as well as his
subsequent settlement in France, and his always writing in their
language, the French claim him as their countryman: an honour
which the Italians are far from conceding to them.
LA GRANGE.
London, Published by
Charles Knight, Pall Mall
East.
The father of Lagrange, luckily perhaps for the fame of his son, was
ruined by some unfortunate speculation. The latter used to say, that
had he possessed fortune, he should probably never have turned his
attention to the science in which he excelled. He was placed at the
College of Turin, and applied himself diligently and with enthusiasm
to classical literature, showing no taste at first for mathematics. In
about a year he began to attend to the geometry of the ancients. A
memoir of Halley in the Philosophical Transactions, on the superiority
of modern analysis, produced consequences of which the author
little dreamed. Lagrange met with it, before his views upon the
subject had settled: and immediately, being then only seventeen
years old, applied himself to the study of the modern mathematics.
Before this change in his studies, according to Delambre[7], after it,
according to others, but certainly while very young, he was elected
professor at the Royal School of Artillery at Turin. We may best
convey some notion of his early proficiency, by stating without detail,
that at the age of twenty-three we find him—the founder of an
Academy of Sciences at Turin, whose volumes yield in interest to
none, and owe that interest principally to his productions,—a
member of the Academy of Sciences at Berlin, an honour obtained
through the medium of Euler, who shortly after announced him to
Frederic of Prussia as the fittest man in Europe to succeed himself,—
and settling, finally, a most intricate question[8] of mathematics,
which had given rise to long discussions between Euler and
D’Alembert, then perhaps the two first mathematicians in Europe. He
had previously extended the method of Euler for the solution of
what are called isoperimetrical problems, and laid the foundation for
the Calculus of Variations, the most decided advance, in our opinion,
which any one has made since the death of Newton.
VOLTAIRE.
London, Published by
Charles Knight, Pall Mall
East.
VOLTAIRE.
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