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Acknowledgements

In preparing the second edition of this book we have again relied heavily on
and benefited greatly from the advice and constructive criticism of numerous
colleagues. We are particularly grateful to Ken Holmes (Max-Planck Institute,
Heidelberg), Lawrence Stern (MIT), Michelle Arkin (Sunesis Pharmaceuticals),
and Watson Fuller (Keele University, UK) for their contributions to, respec-
tively, Chapters 14 and 18, Chapter 15, Chapter 17 and Chapter 18. Stephen
Harrison (Harvard University) and Paul Sigler (Yale University) provided
extensive help and advice on Chapters 8–10, 13 and 16, and Chapters 8–10
and 13, respectively, for which we are especially grateful.
The following, in alphabetical order, have reviewed one or more chap-
ters, correcting our errors of fact or interpretation and helping to ensure they
have the appropriate balance and emphasis: Tom Alber (University of Cali-
fornia, Berkeley), Tom Blundell (Cambridge University, UK), Stephen Burley
(Rockefeller University), Charles Craik (University of California, San Francis-
co), Ken Dill (University of California, San Francisco), Chris Dobson (Oxford
University, UK), Anthony Fink (Unversity of California, Santa Cruz), Robert
Fletterick (University of California, San Francisco), Richard Henderson (LMB,
Cambridge, UK), Werner Kühlbrandt (MPI, Frankfurt), David Parry (Massey
University, New Zealand), Greg Petsko (Brandeis University), and David
Trentham (NIMR, London, UK).
The book depends for its accessibility upon its illustrations and we are
hugely indebted to Nigel Orme, who, as with the first edition, has converted
sketches into lucid figures. Keith Roberts has again advised us on how best
graphically to represent chemical and structural phenomena. Jane Richard-
son (Duke University) has generously produced the Kinemage supplement to
this edition and the book relies upon Richardson-type diagrams throughout
to render the structures discussed comprehensible. We thank our publishers
Garland Publishing, now part of the Taylor and Francis Group, for their sup-
port, and in particular Matthew Day for his enthusiastic editing of the com-
plete manuscript. Miranda Robertson, in her inimitable style, has again
managed the entire project.

vii
Contents

Part I Basic Structural Principles 1 Domains are built from structural motifs 30
1. The Building Blocks 3 Simple motifs combine to form complex motifs 30
Proteins are polypeptide chains 4 Protein structures can be divided into
The genetic code specifies 20 different amino three main classes 31
acid side chains 4 Conclusion 32
Cysteines can form disulfide bridges 5 Selected readings 33
Peptide units are building blocks of protein
structures 8 3. Alpha-Domain Structures 35
Glycine residues can adopt many different Coiled-coil a helices contain a repetitive
conformations 9 heptad amino acid sequence pattern 35
Certain side-chain conformations are The four-helix bundle is a common domain
energetically favorable 10 structure in a proteins 37
Many proteins contain intrinsic metal atoms 11 Alpha-helical domains are sometimes large
Conclusion 12 and complex 39
Selected readings The globin fold is present in myoglobin and
12
hemoglobin 40
2. Motifs of Protein Structure 13 Geometric considerations determine
The interior of proteins is hydrophobic 14 a-helix packing 40
The alpha (a) helix is an important Ridges of one a helix fit into grooves of an
element of secondary structure 14 adjacent helix 40
The a helix has a dipole moment 16 The globin fold has been preserved during
Some amino acids are preferred in evolution 41
a helices 16 The hydrophobic interior is preserved 42
Beta (b) sheets usually have their b strands Helix movements accommodate interior
either parallel or antiparallel 19 side-chain mutations 43
Loop regions are at the surface of Sickle-cell hemoglobin confers resistance
protein molecules 21 to malaria 43
Schematic pictures of proteins highlight Conclusion 45
secondary structure 22 Selected readings 45
Topology diagrams are useful for classification
of protein structures 23 4. Alpha/Beta Structures 47
Secondary structure elements are connected Parallel b strands are arranged in barrels
to form simple motifs 24 or sheets 47
The hairpin b motif occurs frequently in Alpha/beta barrels occur in many different
protein structures 26 enzymes 48
The Greek key motif is found in antiparallel Branched hydrophobic side chains dominate
b sheets 27 the core of a/b barrels 49
The b-a-b motif contains two parallel Pyruvate kinase contains several domains,
b strands 27 one of which is an a/b barrel 51
Protein molecules are organized in a structural Double barrels have occurred by gene
hierarchy 28 fusion 52
Large polypeptide chains fold into several The active site is formed by loops at one
domains 29 end of the a/b barrel 53

ix
 Alpha/beta barrels provide examples of Both single and multiple folding pathways
evolution of new enzyme activities 54 have been observed 93
Leucine-rich motifs form an a/b-horseshoe 55 Enzymes assist formation of proper disulfide
fold bonds during folding 96
Alpha/beta twisted open-sheet structures Isomerization of proline residues can be
contain a helices on both sides of the a rate-limiting step in protein folding 98
b sheet 56 Proteins can fold or unfold inside chaperonins 99
Open b-sheet structures have a variety GroEL is a cylindrical structure with a
of topologies 57 central channel in which newly synthesized
The positions of active sites can be predicted polypeptides bind 100
in a/b structures 57 GroES closes off one end of the GroEL cylinder 102
Tyrosyl-tRNA synthetase has two different The GroEL–GroES complex binds and
domains (a/b + a) 59 releases newly synthesized polypeptides
Carboxypeptidase is an a/b protein with a in an ATP-dependent cycle 102
mixed b sheet 60 The folded state has a flexible structure 104
Arabinose-binding protein has two similar Conformational changes in a protein kinase
a/b domains 62 are important for cell cycle regulation 105
Conclusion 63 Peptide binding to calmodulin induces a
Selected readings 64 large interdomain movement 109
Serpins inhibit serine proteinases with
5. Beta Structures 67 a spring-loaded safety catch mechanism 110
Up-and-down barrels have a simple topology 68 Effector molecules switch allosteric proteins
The retinol-binding protein binds retinol between R and T states 113
inside an up-and-down b barrel 68 X-ray structures explain the allosteric
Amino acid sequence reflects b structure 69 properties of phosphofructokinase 114
The retinol-binding protein belongs to a Conclusion 117
superfamily of protein structures 70 Selected readings 119
Neuraminidase folds into up-and-down b sheets 70
Folding motifs form a propeller-like structure 7. DNA Structures 121
in neuraminidase 71 The DNA double helix is different in A- and
The active site is in the middle of one side of B-DNA 121
the propeller 72 The DNA helix has major and minor grooves 122
Greek key motifs occur frequently in Z-DNA forms a zigzag pattern 123
antiparallel b structures 72 B-DNA is the preferred conformation in vivo 124
The g-crystallin molecule has two domains 74 Specific base sequences can be recognized
The domain structure has a simple topology 74 in B-DNA 124
Two Greek key motifs form the domain 74 Conclusion 125
The two domains have identical topology 75 Selected readings 126
The two domains have similar structures 76
The Greek key motifs in g crystallin are Part 2 Structure, Function, and Engineering 127
evolutionarily related 76 8. DNA Recognition in Procaryotes by
The Greek key motifs can form jelly roll barrels 77 Helix-Turn-Helix Motifs 129
The jelly roll motif is wrapped around a barrel 77 A molecular mechanism for gene control 129
The jelly roll barrel is usually divided into Repressor and Cro proteins operate a procaryotic
two sheets 78 genetic switch region 130
The functional hemagglutinin subunit has two The x-ray structure of the complete lambda
polypeptide chains 79 Cro protein is known 131
The subunit structure is divided into a stem The x-ray structure of the DNA-binding
and a tip 79 domain of the lambda repressor is known 132
The receptor binding site is formed by the Both lambda Cro and repressor proteins
jelly roll domain 80 have a specific DNA-binding motif 133
Hemagglutinin acts as a membrane fusogen 80 Model building predicts Cro–DNA interactions 134
The structure of hemagglutinin is affected Genetic studies agree with the structural model 135
by pH changes 81 The x-ray structure of DNA complexes with
Parallel b-helix domains have a novel fold 84 434 Cro and repressor revealed novel
Conclusion 85 features of protein–DNA interactions 136
Selected readings 87 The structures of 434 Cro and the 434
repressor DNA-binding domain are very
6. Folding and Flexibility 89 similar 137
Globular proteins are only marginally stable 90 The proteins impose precise distortions on
Kinetic factors are important for folding 91 the B-DNA in the complexes 138
Molten globules are intermediates in folding 92 Sequence-specific protein–DNA interactions
Burying hydrophobic side chains is a key event 93 recognize operator regions 138

x
 Protein–DNA backbone interactions determine The finger region of the classic zinc finger
DNA conformation 139 motif interacts with DNA 178
Conformational changes of DNA are Two zinc-containing motifs in the
important for differential binding of repressor glucocorticoid receptor form one
and Cro to different operator sites 140 DNA-binding domain 181
The essence of phage repressor and Cro 141 A dimer of the glucocorticoid receptor binds
DNA binding is regulated by allosteric control 142 to DNA 183
The trp repressor forms a helix-turn-helix motif 142 An a helix in the first zinc motif provides
A conformational change operates a the specific protein–DNA interactions 184
functional switch 142 Three residues in the recognition helix provide
Lac repressor binds to both the major and minor the sequence-specific interactions with DNA 184
grooves inducing a sharp bend in the DNA 143 The retinoid X receptor forms heterodimers
CAP-induced DNA bending could activate that recognize tandem repeats with
transcription 146 variable spacings 185
Conclusion 147 Yeast transcription factor GAL4 contains
Selected readings 148 a binuclear zinc cluster in its DNA-binding
domain 187
9. DNA Recognition by Eucaryotic The zinc cluster regions of GAL4 bind at the
Transcription Factors 151 two ends of the enhancer element 188
Transcription is activated by protein–protein The linker region also contributes to DNA
interactions 152 binding 189
The TATA box-binding protein is ubiquitous 153 DNA-binding site specificity among the C6-zinc
The three-dimensional structures of cluster family of transcription factors is
TBP–TATA box complexes are known 154 achieved by the linker regions 190
A b sheet in TBP forms the DNA-binding site 154 Families of zinc-containing transcription
TBP binds in the minor groove and induces factors bind to DNA in several different ways 191
large structural changes in DNA 155 Leucine zippers provide dimerization
The interaction area between TBP and the interactions for some eucaryotic
TATA box is mainly hydrophobic 157 transcription factors 191
Functional implications of the distortion of The GCN4 basic region leucine zipper binds
DNA by TBP 158 DNA as a dimer of two uninterrupted
159 a helices 193
TFIIA and TFIIB bind to both TBP and DNA
Homeodomain proteins are involved in the GCN4 binds to DNA with both specific and
nonspecific contacts 194
development of many eucaryotic organisms 159
Monomers of homeodomain proteins bind The HLH motif is involved in homodimer
and heterodimer associations 196
to DNA through a helix-turn-helix motif 160
In vivo specificity of homeodomain The a-helical basic region of the b/HLH
motif binds in the major groove of DNA 198
transcription factors depends on interactions
The b/HLH/zip family of transcription factors
with other proteins 162
have both HLH and leucine zipper
POU regions bind to DNA by two tandemly
dimerization motifs 199
oriented helix-turn-helix motifs 164
Max and MyoD recognize the DNA HLH
Much remains to be learnt about the function of
consensus sequence by different specific
homeodomains in vivo 166
protein–DNA interactions 201
Understanding tumorigenic mutations 166 201
Conclusion
The monomeric p53 polypeptide chain is Selected readings 203
divided in three domains 167
The oligomerization domain forms tetramers 167 11. An Example of Enzyme Catalysis:
The DNA-binding domain of p53 is an Serine Proteinases 205
antiparallel b barrel 168 Proteinases form four functional families 205
Two loop regions and one a helix of p53 bind The catalytic properties of enzymes are
to DNA 169 reflected in Km and kcat values 206
Tumorigenic mutations occur mainly in three Enzymes decrease the activation energy of
regions involved in DNA binding 170 chemical reactions 206
Conclusions 172 Serine proteinases cleave peptide bonds by
Selected readings 172 forming tetrahedral transition states 208
Four important structural features are required
10. Specific Transcription Factors Belong 175 for the catalytic action of serine proteinases 209
to a Few Families Convergent evolution has produced two
Several different groups of zinc-containing 176 different serine proteinases with similar
motifs have been observed catalytic mechanisms 210
The classic zinc fingers bind to DNA in tandem 177 The chymotrypsin structure has two antiparallel
along the major groove b-barrel domains 210

xi
 The active site is formed by two loop regions Transmembrane a helices can be predicted
from each domain 211 from amino acid sequences 244
Did the chymotrypsin molecule evolve by Hydrophobicity scales measure the degree
gene duplication? 212 of hydrophobicity of different amino acid
Different side chains in the substrate side chains 245
specificity pocket confer preferential Hydropathy plots identify transmembrane
cleavage 212 helices 245
Engineered mutations in the substrate Reaction center hydropathy plots agree with
specificity pocket change the rate of catalysis 213 crystal structural data 246
The Asp 189-Lys mutation in trypsin causes Membrane lipids have no specific interaction
unexpected changes in substrate specificity 215 with protein transmembrane a helices 246
The structure of the serine proteinase subtilisin Conclusion 247
is of the a/b type 215 Selected readings 248
The active sites of subtilisin and chymotrypsin
are similar 216 13. Signal Transduction 251
A structural anomaly in subtilisin has G proteins are molecular amplifiers 252
functional consequences 217 Ras proteins and the catalytic domain
Transition-state stabilization in subtilisin is of Ga have similar three-dimensional 254
dissected by protein engineering 217 structures
Catalysis occurs without a catalytic triad 217 Ga is activated by conformational changes of
Substrate molecules provide catalytic groups three switch regions 257
in substrate-assisted catalysis 218 GTPases hydrolyze GTP through nucleophilic
Conclusion 219 attack by a water molecule 259
Selected readings 220 The Gb subunit has a seven-blade propeller fold,
built up from seven WD repeat units 261
12. Membrane Proteins 223 The GTPase domain of Ga binds to Gb in the
Membrane proteins are difficult to crystallize 224 heterotrimeric Gabg complex 263
Novel crystallization methods are being Phosducin regulates light adaptation in
developed 224 265
retinal rods
Two-dimensional crystals of membrane Phosducin binding to Gbg blocks binding of Ga 265
proteins can be studied by electron
The human growth hormone induces
microscopy 225
dimerization of its cognate receptor 267
Bacteriorhodopsin contains seven
Dimerization of the growth hormone receptor
transmembrane a helices 226
is a sequential process 268
Bacteriorhodopsin is a light-driven proton
The growth hormone also binds to the
pump 227
prolactin receptor 269
Porins form transmembrane channels by
Tyrosine kinase receptors are important
b strands 228
enzyme-linked receptors 270
Porin channels are made by up and down
Small protein modules form adaptors for a
b barrels 229
signaling network 272
Each porin molecule has three channels 230
SH2 domains bind to phosphotyrosine-
Ion channels combine ion selectivity with
containing regions of target molecules 273
high levels of ion conductance 232
SH3 domains bind to proline-rich regions of
The K+ channel is a tetrameric molecule with target molecules 274
one ion pore in the interface between the Src tyrosine kinases comprise SH2 and SH3
four subunits 232
domains in addition to a tyrosine kinase 275
The ion pore has a narrow ion selectivity filter 233
The two domains of the kinase in the
The bacterial photosynthetic reaction center inactive state are held in a closed
is built up from four different polypeptide conformation by assembly of the
chains and many pigments 234
regulatory domains 277
The L, M, and H subunits have transmembrane Conclusion 278
a helices 236
Selected readings 280
The photosynthetic pigments are bound to the
L and M subunits 237
14. Fibrous Proteins 283
Reaction centers convert light energy into Collagen is a superhelix formed by three
electrical energy by electron flow through parallel, very extended left-handed helices 284
the membrane 239
Coiled coils are frequently used to form
Antenna pigment proteins assemble into oligomers of fibrous and globular proteins 286
multimeric light-harvesting particles 240
Amyloid fibrils are suggested to be built up
Chlorophyll molecules form circular rings from continuous b sheet helices 288
in the light-harvesting complex LH2 241
Spider silk is nature’s high-performance fiber 289
The reaction center is surrounded by a ring of 16 Muscle fibers contain myosin and actin which
antenna proteins of the light-harvesting slide against each other during muscle
complex LH1 242
contraction 290

xii
 Myosin heads form cross-bridges between the Complex spherical viruses have more than one
actin and myosin filaments 291 polypeptide chain in the asymmetric unit 329
Time-resolved x-ray diffraction of frog muscle Structural versatility gives quasi-equivalent
confirmed movement of the cross-bridges 292 packing in T = 3 plant viruses 331
Structures of actin and myosin have been The protein subunits recognize specific parts
determined 293 of the RNA inside the shell 332
The structure of myosin supports the swinging The protein capsid of picornaviruses contains
cross-bridge hypothesis 295 four polypeptide chains 333
The role of ATP in muscular contraction has There are four different structural proteins in
parallels to the role of GTP in G-protein picornaviruses 334
activation 296 The arrangement of subunits in the shell of
Conclusion 297 picornaviruses is similar to that of T = 3
Selected readings 298 plant viruses 334
The coat proteins of many different spherical
15. Recognition of Foreign Molecules by the plant and animal viruses have similar
Immune System 299 jelly roll barrel structures, indicating an
The polypeptide chains of antibodies are evolutionary relationship 335
divided into domains 300 Drugs against the common cold may be
Antibody diversity is generated by several designed from the structure of rhinovirus 337
different mechanisms 302 Bacteriophage MS2 has a different subunit
All immunoglobulin domains have similar structure 339
three-dimensional structures 303 A dimer of MS2 subunits recognizes an RNA
The immunoglobulin fold is best described as packaging signal 339
two antiparallel b sheets packed tightly The core protein of alphavirus has a
against each other 304 chymotrypsin-like fold 340
The hypervariable regions are clustered SV40 and polyomavirus shells are constructed
in loop regions at one end of the from pentamers of the major coat protein
variable domain 305 with nonequivalent packing but largely
The antigen-binding site is formed by close equivalent interactions 341
association of the hypervariable regions Conclusion 343
from both heavy and light chains 306 Selected readings 344
The antigen-binding site binds haptens in
crevices and protein antigens on large 17. Prediction, Engineering, and Design of
flat surfaces 308 Protein Structures 347
The CDR loops assume only a limited range Homologous proteins have similar structure
of conformations, except for the heavy and function 348
chain CDR3 311 Homologous proteins have conserved structural
An IgG molecule has several degrees of cores and variable loop regions 349
conformational flexibility 312 Knowledge of secondary structure is necessary
Structures of MHC molecules have provided for prediction of tertiary structure 350
insights into the molecular mechanisms Prediction methods for secondary structure
of T-cell activation 312 benefit from multiple alignment of
MHC molecules are composed of antigen- homologous proteins 351
binding and immunoglobulin-like domains 313 Many different amino acid sequences give
Recognition of antigen is different in MHC similar three-dimensional structures 352
molecules compared with immunoglobulins 314 Prediction of protein structure from sequence
Peptides are bound differently by class I and is an unsolved problem 352
class II MHC molecules 315 Threading methods can assign amino acid
T-cell receptors have variable and constant sequences to known three-dimensional folds 353
immunoglobulin domains and Proteins can be made more stable by
hypervariable regions 316 engineering 354
MHC–peptide complexes are the ligands for Disulfide bridges increase protein stability 355
T-cell receptors 318 Glycine and proline have opposite effects on
Many cell-surface receptors contain stability 356
immunoglobulin-like domains. 318 Stabilizing the dipoles of a helices increases
Conclusion 320 stability 357
Selected readings 321 Mutants that fill cavities in hydrophobic cores
do not stabilize T4 lysozyme 358
16. The Structure of Spherical Viruses 325 Proteins can be engineered by combinatorial
The protein shells of spherical viruses have methods 358
icosahedral symmetry 327 Phage display links the protein library
The icosahedron has high symmetry 327 to DNA 359
The simplest virus has a shell of 60 protein Affinity and specificity of proteinase inhibitors
subunits 328 can be optimized by phage display 361

xiii
 Structural scaffolds can be reduced in size Building a model involves subjective
while function is retained 363 interpretation of the data 381
Phage display of random peptide libraries Errors in the initial model are removed
identified agonists of erythropoetin receptor 364 by refinement 383
DNA shuffling allows accelerated evolution Recent technological advances have greatly
of genes 365 influenced protein crystallography 383
Protein structures can be designed from first X-ray diffraction can be used to study the
principles 367 structure of fibers as well as crystals 384
A b structure has been converted to an a structure The structure of biopolymers can be studied
by changing only half of the sequence 368 using fiber diffraction 386
Conclusion 370 NMR methods use the magnetic properties of
Selected readings 371 atomic nuclei 387
Two-dimensional NMR spectra of proteins are
18. Determination of Protein Structures 373 interpreted by the method of sequential
Several different techniques are used to study assignment 389
the structure of protein molecules 373 Distance constraints are used to derive possible
Protein crystals are difficult to grow 374 structures of a protein molecule 390
X-ray sources are either monochromatic or Biochemical studies and molecular
polychromatic 376 structure give complementary functional
X-ray data are recorded either on image plates information 391
or by electronic detectors 377 Conclusion 391
The rules for diffraction are given by Bragg’s law 378 Selected readings 392
Phase determination is the major
crystallographic problem 379 Protein Structure on the World Wide Web 393
Phase information can also be obtained by
Multiwavelength Anomalous Diffraction
experiments 381

xiv
Basic Part 1
Structural
Principles

1
This page is intentionally left blank.
The Building Blocks
1

Recombinant DNA techniques have provided tools for the rapid determination
of DNA sequences and, by inference, the amino acid sequences of proteins
from structural genes. The number of such sequences is now increasing
almost exponentially, but by themselves these sequences tell little more
about the biology of the system than a New York City telephone directory
tells about the function and marvels of that city.
The proteins we observe in nature have evolved, through selective pres-
sure, to perform specific functions. The functional properties of proteins Figure 1.1 The amino acid sequence of a
depend upon their three-dimensional structures. The three-dimensional protein’s polypeptide chain is called its
primary structure. Different regions of the
structure arises because particular sequences of amino acids in polypeptide
sequence form local regular secondary
chains fold to generate, from linear chains, compact domains with specific structures, such as alpha (a) helices or beta (b)
three-dimensional structures (Figure 1.1). The folded domains can serve as strands. The tertiary structure is formed by
modules for building up large assemblies such as virus particles or muscle packing such structural elements into one or
fibers, or they can provide specific catalytic or binding sites, as found in several compact globular units called domains.
enzymes or proteins that carry oxygen or that regulate the function of DNA. The final protein may contain several
polypeptide chains arranged in a quaternary
To understand the biological function of proteins we would therefore
structure. By formation of such tertiary and
like to be able to deduce or predict the three-dimensional structure from the quaternary structure amino acids far apart in
amino acid sequence. This we cannot do. In spite of considerable efforts over the sequence are brought close together in
the past 25 years, this folding problem is still unsolved and remains one of three dimensions to form a functional region,
the most basic intellectual challenges in molecular biology. an active site.

Primary Secondary Tertiary Quarternary

F T P A
V C C
L F A H C
D
K F L A N N
S N N
V T S V N
L
C
T S K Y C

3
 Protein folding remains a problem because there are 20 different amino
acids that can be combined into many more different proteins than there are
atoms in the known universe. In addition there is a vast number of ways in
which similar structural domains can be generated in proteins by different
amino acid sequences. By contrast, the structure of DNA, made up of only
four different nucleotide building blocks that occur in two pairs, is relatively
simple, regular, and predictable.
Since the three-dimensional structures of individual proteins cannot be
predicted, they must instead be determined experimentally by x-ray crystal-
lography, electron crystallography or nuclear magnetic resonance (NMR)
techniques. Over the past 30 years the structures of more than 6000 proteins
have been solved, and the sequences of more than 500,000 have been deter-
mined. This has generated a body of information from which a set of basic
principles of protein structure has emerged. These principles make it easier
for us to understand how protein structure is generated, to identify common
structural themes, to relate structure to function, and to see fundamental
relationships between different proteins. The science of protein structure is at
the stage of taxonomy where we can begin to discern patterns and motifs
among the relatively small number of proteins whose three-dimensional
structure is known. Figure 1.2 Proteins are built up by amino
The first six chapters of this book deal with the basic principles of pro- acids that are linked by peptide bonds to form
tein structure as we understand them today, and examples of the different a polypeptide chain. (a) Schematic diagram of
an amino acid, illustrating the nomenclature
major classes of protein structures are presented. Chapter 7 contains a brief
used in this book. A central carbon atom (Ca)
discussion on DNA structures with emphasis on recognition by proteins of is attached to an amino group (NH2), a
specific nucleotide sequences. The remaining chapters illustrate how during carboxyl group (COOH), a hydrogen atom (H),
evolution different structural solutions have been selected to fulfill particular and a side chain (R). (b) In a polypeptide chain
functions. the carboxyl group of amino acid n has formed
a peptide bond, C–N, to the amino group
of amino acid n + 1. One water molecule is
Proteins are polypeptide chains eliminated in this process. The repeating units,
which are called residues, are divided into
All of the 20 amino acids have in common a central carbon atom (Ca) to main-chain atoms and side chains. The
which are attached a hydrogen atom, an amino group (NH2), and a carboxyl main-chain part, which is identical in all
group (COOH) (Figure 1.2a). What distinguishes one amino acid from residues, contains a central Ca atom attached
another is the side chain attached to the Ca through its fourth valence. There to an NH group, a C¢=O group, and an H atom.
The side chain R, which is different for
are 20 different side chains specified by the genetic code; others occur, in rare
different residues, is bound to the Ca atom.
cases, as the products of enzymatic modifications after translation.
Amino acids are joined end-to-end during protein synthesis by the for-
(a) side chain
mation of peptide bonds when the carboxyl group of one amino acid con-
denses with the amino group of the next to eliminate water (Figure 1.2b).
R
This process is repeated as the chain elongates. One consequence is that the
amino group of the first amino acid of a polypeptide chain and the carboxyl
group of the last amino acid remain intact, and the chain is said to extend H OH
Cα
from its amino terminus to its carboxy terminus. The formation of a succes- N C¢
sion of peptide bonds generates a “main chain,” or “backbone,” from which
project the various side chains. H
H O
The main-chain atoms are a carbon atom Ca to which the side chain is
attached, an NH group bound to Ca, and a carbonyl group C¢=O, where the amino group carboxyl group
carbon atom C¢ is attached to Ca. These units, or residues, are linked into a
polypeptide by a peptide bond between the C¢ atom of one residue and the
nitrogen atom of the next (see Figure 1.2b). The basic repeating unit along (b) peptide bond
the main chain from a biochemical or genetic viewpoint is thus
(NH–CaH–C¢=O), which is the residue of the common parts of amino acids
after peptide bonds have been formed (see Figure 1.2b).
R1 H H O

The genetic code specifies 20 different amino acid side chains N Cα

C¢
N Cα C¢
The 20 different side chains that occur in proteins are shown in Panel 1.1 (pp. H R2
6–7). Their names are abbreviated with both a three-letter and a one-letter H O
code, which are also given in the panel. The one-letter codes are worth mem-
orizing, as they are widely used in the literature. A mnemonic device for link- n n+1
ing the one-letter code to the names of the amino acids is given in Panel 1.1.

4
 The amino acids are usually divided into three different classes defined R R
by the chemical nature of the side chain. The first class comprises those with
strictly hydrophobic side chains: Ala (A), Val (V), Leu (L), Ile (I), Phe (F), Pro
(P), and Met (M). The four charged residues, Asp (D), Glu (E), Lys (K), and Arg Cα Cα
(R), form the second class. The third class comprises those with polar side CO H N N H CO
chains: Ser (S), Thr (T), Cys (C), Asn (N), Gln (Q), His (H), Tyr (Y), and Trp L-form D-form
(W). The amino acid glycine (G), which has only a hydrogen atom as a side
chain and so is the simplest of the 20 amino acids, has special properties and Figure 1.3 The “handedness” of amino acids.
is usually considered either to form a fourth class or to belong to the first Looking down the H–Ca bond from the
hydrogen atom, the L-form has CO, R, and
class.
N substituents from Ca going in a clockwise
The four groups attached to the Ca atom are chemically different for all direction. There is a mnemonic to remember
the amino acids except glycine, where two H atoms bind to Ca. All amino this; for the L-form the groups read CORN in
acids except glycine are thus chiral molecules that can exist in two different clockwise direction.
forms with different “hands,” L- or D-form (Figure 1.3).
Biological systems depend on specific detailed recognition of molecules
that distinguish between chiral forms. The translation machinery for protein
synthesis has evolved to utilize only one of the chiral forms of amino acids,
the L-form. All amino acids that occur in proteins therefore have the L-form.
There is, however, no obvious reason why the L-form was chosen during evo-
lution and not the D-form.

Cysteines can form disulfide bridges

Two cysteine residues in different parts of the polypeptide chain but adjacent
in the three-dimensional structure of a protein can be oxidized to form a
disulfide bridge (Figure 1.4). The disulfide is usually the end product of air
oxidation according to the following reaction scheme:
2 –CH2SH + © O2  –CH2–S–S–CH2 + H2O
This reaction requires an oxidative environment, and such disulfide bridges
are usually not found in intracellular proteins, which spend their lifetime in
an essentially reductive environment. Disulfide bridges do, however, occur
quite frequently among extracellular proteins that are secreted from cells,
and in eucaryotes, formation of these bridges occurs within the lumen of the
endoplasmic reticulum, the first compartment of the secretory pathway.

cysteine cysteine

S
H
+
H
S

oxidation reduction

Figure 1.4 The disulfide is usually the end

product of air oxidation according to the
following schematic reaction scheme:
S 2 –CH2SH + © O2  –CH2–S–S–CH2 + H2O
Disulfide bonds form between the side chains
of two cysteine residues. Two SH groups from
cysteine residues, which may be in different
disulfide bond S parts of the amino acid sequence but adjacent
in the three-dimensional structure, are
oxidized to form one S–S (disulfide) group.

5
 (a) Hydrophobic amino acids
H
C
HC CH
CH3 H 3C CH3 HC CH
– CH C
H3N+ C COO
CH2
H
A Ala, Alanine V Val, Valine F Phe, Phenylalanine

H3C CH3
CH3
CH
H2C CH3
CH CH 2

I Ile, Isoleucine L Leu, Leucine

(b) Charged amino acids +

NH3
O O–
CH 2
O O– C
CH 2
C CH2
CH 2
CH2 CH2
CH 2

D Asp, Aspartic acid E Glu, Glutamic acid K Lys, Lysine

(c) Polar amino acids OH

OH HO CH3
CH
CH2 CH2

S Ser, Serine T Thr, Threonine Y Tyr, Tyrosine

O NH2
O NH2 C
SH C
CH2
CH2 CH2
CH2

C Cys, Cysteine N Asn, Asparagine Q Gln, Glutamine

6
 Panel 1.1 The 20 different amino acids that
CH3 occur in proteins. Only side chains are shown,
except for the first amino acid, alanine, where
S all atoms are shown. The bond from the side
chain to Ca is red. A ball-and-stick model, the
CH2 CH2 CH2 chemical formula, the full name, and the
three-letter and one-letter codes are given for
CH2 C αH CH2 each amino acid.
N There are some easy ways of remembering
the one-letter code for amino acids. If only one
amino acid begins with a certain letter, that
P Pro, Proline M Met, Methionine letter is used:

C = Cys = Cysteine
H = His = Histidine
I = Ile = Isoleucine
M = Met = Methionine
S = Ser = Serine
V = Val = Valine

If more than one amino acid begins with a

certain letter, that letter is assigned to the
most commonly occurring amino acid:

A = Ala = Alanine
G = Gly = Glycine
L = Leu = Leucine
NH2 P = Pro = Proline
+ T = Thr = Threonine
C=NH2
Some of the others are phonetically suggestive:
NH
F = Phe = Phenylalanine (“Fenylalanine”)
CH 2 R = Arg = Arginine (“aRginine”)
Y = Tyr = Tyrosine (“tYrosine”)
CH 2 W = Trp = Tryptophan (double ring in
the molecule)
CH 2
R Arg, Arginine In other cases a letter close to the initial is
used. Amides have letters from the middle of
the alphabet. The smaller molecules (D, N, B)
are earlier in the alphabet than the larger ones
(E, Q, Z).

D = Asp = Aspartic acid (near A)

HN N = Asn = Asparagine (contains N)
+ B = Asx = Either of D or N
NH E = Glu = Glutamic acid (near G)
Q = Gln = Glutamine (“Q-tamine”)
CH2 Z = Glx = Either of E or Q
K = Lys = Lysine (near L)
X = X = Undetermined amino acid

H His, Histidine

H (d) Glycine
N

H
CH2

G Gly, Glycine
W Trp, Tryptophan

7
 Figure 1.5 Part of a polypeptide chain that
O R2 H is divided into peptide units, represented
as blocks in the diagram. Each peptide unit
H C¢ Cα H contains the Ca atom and the C¢=O group of
N
N residue n as well as the NH group and the
Cα C¢ Ca atom of residue n + 1. Each such unit is a
H Cα
planar, rigid group with known bond distances
H O and bond angles. R1, R2, and R3 are the side
R1 R3 chains attached to the Ca atoms that link the
peptide units in the polypeptide chain. The
peptide bond
peptide group is planar because the additional
electron pair of the C=O bond is delocalized
over the peptide group such that rotation
around the C–N bond is prevented by an
Disulfide bridges stabilize three-dimensional structure. In some proteins energy barrier.
these bridges hold together different polypeptide chains; for example, the A
and B chains of insulin are linked by two disulfide bridges between the
chains. More frequently intramolecular disulfide bridges stabilize the folding
of a single polypeptide chain, making the protein less susceptible to degra-
dation. There are many examples of this, including proteins with short
polypeptide chains, such as snake venom toxins and protease inhibitors, that
need additional stabilizing factors to produce a stable fold. Much effort is
currently spent on introducing extra intramolecular disulfide bridges into
enzymes by site-directed mutagenesis in order to make them more thermo-
stable and hence more useful for industrial applications as catalysts, as
described in Chapter 17.

Peptide units are building blocks of protein structures psi3

Figure 1.2 shows one way of dividing a polypeptide chain, the biochemist’s
way. There is, however, a different way to divide the main chain into repeat-
ing units that is preferable when we want to describe the structural proper-
phi3
ties of proteins. For this purpose it is more useful to divide the polypeptide
chain into peptide units that go from one Ca atom to the next Ca atom (see
Figure 1.5). Each Ca atom, except the first and the last, thus belongs to two
such units. The reason for dividing the chain in this way is that all the atoms
in such a unit are fixed in a plane with the bond lengths and bond angles
very nearly the same in all units in all proteins. Note that the peptide units
of the main chain do not involve the different side chains (Figure 1.5). We psi2
will use both of these alternative descriptions of polypeptide chains—the bio-
chemical and the structural—and discuss proteins in terms of the sequence
of different amino acids and the sequence of planar peptide units.
Since the peptide units are effectively rigid groups that are linked into a phi2
chain by covalent bonds at the Ca atoms, the only degrees of freedom they
have are rotations around these bonds. Each unit can rotate around two such
bonds: the Ca–C¢ and the N–Ca bonds (Figure 1.6). By convention the angle H
of rotation around the N–Ca bond is called phi (f f) and the angle around the
Ca–C¢ bond from the same Ca atom is called psi (y y). psi1 Ca
In this way each amino acid residue is associated with two conforma-
tional angles f and y. Since these are the only degrees of freedom, the con- phi1 R
formation of the whole main chain of the polypeptide is completely deter-
mined when the f and y angles for each amino acid are defined with high
accuracy. H
N
C¢
peptide plane BT2 O
Figure 1.6 Diagram showing a polypeptide chain where the main-chain 1.4/1.4
atoms are represented as rigid peptide units, linked through the Ca
atoms. Each unit has two degrees of freedom; it can rotate around two
bonds, its Ca–C¢ bond and its N–Ca bond. The angle of rotation around
the N–Ca bond is called phi (f) and that around the Ca–C¢ bond is
called psi (y). The conformation of the main-chain atoms is therefore
determined by the values of these two angles for each amino acid. Cα

8
Glycine residues can adopt many different conformations
Most combinations of f and y angles for an amino acid are not allowed
because of steric collisions between the side chains and main chain. It is rea-
sonably straightforward to calculate those combinations that are allowed.
Since the D- and L-forms of the amino acids have their side chain oriented
differently with respect to the CO group, they have different allowed f and
y angles. Proteins built from D-amino acids would thus be expected to have
different conformations from those found in nature that are exclusively
made of L-amino acids. Since the L- and D-forms of each amino acid are mir-
ror images of one another, would a protein made exclusively of D-form
residues produce a structure that is the mirror image of the natural protein?
Stephen Kent and his colleagues at the Scripps Institute chemically synthe-
sized both the L- and the D-forms of HIV-1 protease. The D-enzyme proved
indeed to be the mirror image of the L-enzyme. Furthermore the D-enzyme
and L-enzyme had reciprocal chiral specificity on peptide substrates, the
D-enzyme only recognizing and cutting peptides made of D-amino acids. Per-
haps the choice of the L-form at the outset of the evolution of life on earth
was random and irrevocable.
The angle pairs f and y are usually plotted against each other in a
diagram called a Ramachandran plot after the Indian biophysicist
G.N. Ramachandran who first made calculations of sterically allowed regions.
Figure 1.7 shows the results of such calculations and also a plot for all amino
Figure 1.7 Ramachandran plots showing
(a) allowed combinations of the conformational
angles phi and psi defined in Figure 1.6. Since
+180 phi (f) and psi (y) refer to rotations of two
rigid peptide units around the same Ca atom,
β most combinations produce steric collisions
either between atoms in different peptide
groups or between a peptide unit and the side
L chain attached to Ca. These combinations are
therefore not allowed. (a) Colored areas show
sterically allowed regions. The areas labeled
psi 0 a, b, and L correspond approximately to
conformational angles found for the usual
α
right-handed a helices, b strands, and
left-handed a helices, respectively. (b)
Observed values for all residue types except
glycine. Each point represents f and y values
for an amino acid residue in a well-refined
–180 x-ray structure to high resolution. (c) Observed
values for glycine. Notice that the values
–180 0 phi +180 include combinations of f and y that are not
allowed for other amino acids. (From J.
Richardson, Adv. Prot. Chem. 34: 174–175,
1981.)
(b) (c)
+180 +180

psi 0 psi 0

–180 –180
–180 0 phi +180 –180 0 phi +180

9
acids except glycine from a number of accurately determined protein struc-
tures. It is apparent that the observed values are clustered in the sterically
allowed regions. There is one important exception. Glycine, with only a
hydrogen atom as a side chain, can adopt a much wider range of conforma-
tions than the other residues, as seen in Figure 1.7c. Glycine thus plays a
structurally very important role; it allows unusual main-chain conformations
in proteins. This is one of the main reasons why a high proportion of glycine
residues are conserved among homologous protein sequences.
Regions in the Ramachandran plot are named after the conformation
that results in a peptide if the corresponding f and y angles are repeated in
successive amino acids along the chain. The major allowed regions in Figure
1.7a are the right-handed a-helical cluster in the lower left quadrant (see
Chapter 3); the broad region of extended b strands of both parallel and
antiparallel b structures (see Chapter 4) in the upper left quadrant; and the
small, sparsely populated left-handed a-helical region in the upper right
quadrant. Left-handed a helices are usually found in loop regions or in small
single-turn a helices.

Certain side-chain conformations are energetically favorable

Any side chain longer than that of alanine can in principle have several dif-
ferent conformations because of rotations around the bonds between the
side-chain carbon atoms. It is a general rule in chemistry that the most ener-
getically favored arrangements for two tetrahedrally coordinated carbon
atoms are the “staggered” conformations, in which the substituents of one
carbon atom are between those of the other when viewed along the axis of
rotation, as illustrated in Figure 1.8. For each such carbon atom in a side
chain, there are three possible staggered conformations, which are related by
a three step, 120˚ rotation around the carbon–carbon bond. These three con-
formations are indistinguishable for alanine, since all three substituents on
the side-chain carbon atom Cb are the same.
For valine, however, the three staggered conformations are energetically dif-
ferent because the substituents on Cb are different; two of them are methyl
groups and the other is a hydrogen atom (see Figure 1.8b–d). It is energetically
most favorable to have the two methyl groups close to the small hydrogen atom
bound to Ca, as shown in Figure 1.8b, and therefore this is the
conformation most frequently found in proteins. An analysis of accurately
determined protein structures has shown that most side chains have one or a
few conformations that occur more frequently than the other possible staggered

(a) not staggered staggered

Figure 1.8 The staggered conformations are

the most energetically favored conformations
of two tetrahedrally coordinated carbon atoms.
(a) A view along the C–C bond in ethane
(CH3CH3) showing how the two carbon atoms
can rotate so that their hydrogen atoms are
either not staggered (aligned) or staggered.Three
(b) CH3 CH3 (c) CH3 (d) CH3 indistinguishable staggered conformations are
H H
obtained by a rotation of 120˚ around the C–C
H H H bond. (b–d) Similar views as in (a) of valine.
The three staggered conformations are different
for valine because the three groups attached
CH3 CH3 to Cb are different. The first staggered
H conformation (b) is less crowded and
N energetically most favored because the two
N N
O O O methyl groups bound to Cb are both close to
H H H the small H atom bound to Ca.

10
 (a) (b)
Tyr (free radical)
Cys
coenzyme S Cys
O O
Glu S
μ-oxo bridge
H2O H2O O Zn
O O O
O 2–
Glu His
Fe Fe N
Asp O
O –
O O N
N N alcohol

His Glu His

N N

conformations. These are called rotamers. Today, collections of these favored Figure 1.9 Examples of functionally important
conformations, or rotamer libraries, are a standard tool in computer programs intrinsic metal atoms in proteins. (a) The
used for modeling protein structures. di-iron center of the enzyme ribonucleotide
reductase. Two iron atoms form a redox center
that produces a free radical in a nearby tyrosine
Many proteins contain intrinsic metal atoms side chain. The iron atoms are bridged by a
glutamic acid residue and a negatively charged
The side chains of the 20 different amino acids listed in Panel 1.1 (pp. 6–7) oxygen atom called a m-oxo bridge. The
have very different chemical properties and are utilized for a wide variety of coordination of the iron atoms is completed
by histidine, aspartic acid, and glutamic acid
biological functions. However, their chemical versatility is not unlimited,
side chains as well as water molecules. (b) The
and for some functions metal atoms are more suitable and more efficient. catalytically active zinc atom in the enzyme
Electron-transfer reactions are an important example. Fortunately the side alcohol dehydrogenase. The zinc atom is
chains of histidine, cysteine, aspartic acid, and glutamic acid are excellent coordinated to the protein by one histidine
metal ligands, and a fairly large number of proteins have recruited metal and two cysteine side chains. During catalysis
atoms as intrinsic parts of their structures; among the frequently used metals zinc binds an alcohol molecule in a suitable
are iron, zinc, magnesium, and calcium. Several metallo proteins are dis- position for hydride transfer to the coenzyme
moiety, a nicotinamide. [(a) Adapted from
cussed in detail in later chapters and it suffices here to mention briefly a few
P. Nordlund et al., Nature 345: 593–598, 1990.]
examples of iron and zinc proteins.
The most conspicuous use of iron in biological systems is in our blood,
where the erythrocytes are filled with the oxygen-binding protein hemoglo-
bin. The red color of blood is due to the iron atom bound to the heme group
in hemoglobin. Similar heme-bound iron atoms are present in a number
of proteins involved in electron-transfer reactions, notably cytochromes. A
chemically more sophisticated use of iron is found in an enzyme, ribo-
nucleotide reductase, that catalyzes the conversion of ribonucleotides to
deoxyribonucleotides, an important step in the synthesis of the building
blocks of DNA.
Ribonucleotide reductase from Escherichia coli and mammals contains a
di-iron center (Figure 1.9a) that reacts with oxygen and oxidizes a nearby
tyrosine side chain, producing a tyrosyl free radical that is essential for the
catalysis. The two iron atoms in this iron center are close to each other and
bridged by the oxygen atoms of a glutamic acid side chain as well as by an
oxygen ion called a m-oxo bridge. Glutamic acid, aspartic acid, and histidine
side chains from the protein, as well as water molecules, complete the co-
ordination sphere of the octahedrally coordinated iron atom.
Zinc is used to stabilize the DNA-binding regions of a class of transcrip-
tion factors called zinc fingers, which are discussed in Chapter 10. Zinc ions
also participate directly in catalytic reactions in many different enzymes by
binding substrate molecules and providing a positive charge that influences
the electronic arrangement of the substrate and thereby facilitates the cat-
alytic reaction. One such example is found in the enzyme alcohol dehydro-
genase, which in yeast produces alcohol during fermentation and in our
livers detoxifies the alcohol we have consumed by oxidizing it. The enzyme
provides a scaffold containing three zinc ligands—one histidine and two cys-
teine side chains—which sequester a zinc atom so that it binds alcohol as a
fourth ligand in a tetrahedral coordination (Figure 1.9b).

11
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Title: Nightmare Planet

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*** START OF THE PROJECT GUTENBERG EBOOK NIGHTMARE

PLANET ***
Nightmare Planet
 by MURRAY LEINSTER
(Illustrations by Tom O'Reilly)
[Transcriber's Note: This etext was produced from Science Fiction
Plus June 1953. Extensive research did not uncover any evidence
that the U.S. copyright on this publication was renewed.]

In science-fiction, as in all categories of

fiction, there are stories that are so
outstanding from the standpoint of
characterization, concept, and
background development that they
remain popular for decades. Two such
stories were Murray Leinster's The Mad
Planet and Red Dust. Originally
published in 1923, they have been
reprinted frequently both here and
abroad. They are now scheduled for
book publication. Especially for this
magazine, Murray Leinster has written
the final story in the series. It is not
necessary to have read the previous
stories to enjoy this one. Once again, Burl experiences magnificent
adventures against a colorful background, but to the whole the
author has added philosophical and psychological observations that
give this story a flavor seldom achieved in science-fiction.
Under his real name of Will Fitzgerald Jenkins, the author has sold to
The Saturday Evening Post, Colliers', Today's Woman, in fact every
important publication in America. He has had over 1200 stories
published, 15 books and 35 science-fiction stories anthologized. His
writing earned him a listing in Who's Who in America.

The Directory-ship Tethys made the first landing on the planet,

L21612. It was a goodly world, with an ample atmosphere and many
seas, which the nearby sun warmed so lavishly that a perpetual
cloud-bank hid them and all the solid ground from view. It had
mountains and islands and high plateaus. It had day and night and
rain. It had an equable climate, rather on the tropical side. But it
possessed no life.
No animals roamed its solid surface. No vegetation grew from its
rocks. Not even bacteria struggled with the stones to turn them into
soil. No living thing, however small, swam in its oceans. It was one
of that disappointing vast majority of otherwise admirable worlds
which was unsuited for colonization solely because it had not been
colonized before. It could be used for biological experiments in a
completely germ-free environment, or ships could land upon it for
water and supplies of air. The water was pure and the air
breathable, but it had no other present utility. Such was the case
with an overwhelming number of Earth-type planets when first
discovered in the exploration of the galaxy. Life simply hadn't started
there.
So the ship which first landed upon it made due note for the Galactic
Directory and went away, and no other ship came near the planet
for eight hundred years.
But nearly a millennium later, the Seed-Ship Orana arrived. It landed
and carefully seeded the useless world. It circled endlessly above the
clouds, dribbling out a fine dust comprised of the spores of every
conceivable microorganism that could break down rock to powder
and turn the powder to organic matter. It also seeded with moulds
and fungi and lichens, and everything that could turn powdery
primitive soil into stuff on which higher forms of life could grow. The
Orana seeded the seas with plankton. Then it, too, went away.
Centuries passed. Then the Ecological Preparation Ship Ludred swam
to the planet from space. It was a gigantic ship of highly improbable
construction and purpose. It found the previous seeding successful.
Now there was soil which swarmed with minute living things. There
were fungi which throve monstrously. The seas stank of teeming
minuscule life-forms. There were even some novelties on land,
developed by strictly local conditions. There were, for example,
paramecium as big as grapes, and yeasts had increased in size so
that they bore flowers visible to the naked eye. The life on the
planet was not aboriginal, though. It had all been planted by the
seed-ship of centuries before.
The Ludred released insects, it dumped fish into the seas. It
scattered plant-seeds over the continents. It treated the planet to a
sort of Russell's Mixture of living things. The real Russell's Mixture is
that blend of simple elements in the proportions found in suns. This
was a blend of living creatures, of whom some should certainly
survive by consuming the now habituated flora, and others which
should survive by preying on the first. The planet was stocked, in
effect, with everything it could be hoped might live there.
But at the time of the Ludred's visit of course no creature needing
parental care had any chance of survival. Everything had to be able
to care for itself the instant it burst its egg. So there were no birds
or mammals. Trees and plants of divers sorts, and fish and
crustaceans and insects could be planted. Nothing else.
The Ludred swam away through emptiness.
There should have been another planting, centuries later still, but it
was never made. When the Ecological Preparation Service was
moved to Algol IV, a file was upset. The cards in it were picked up
and replaced, but one was missed. So that planet was forgotten. It
circled its sun in emptiness. Cloud-banks covered it from pole to
pole. There were hazy markings in certain places, where high
plateaus penetrated the clouds. But from space the planet was
featureless. Seen from afar, it was merely a round white ball—white
from its cloud-banks and nothing else.
But on its surface, in its lowlands it was nightmare.
Especially was it nightmare—after some centuries—for the
descendants of the human beings from the space-liner Icarus,
wrecked there some forty-odd generations ago. Naturally, nobody
anywhere else thought of the Icarus any more. It was not even
remembered by the descendants of its human cargo, who now
inhabited the planet. The wreckage of the ship was long since
hidden under the seething, furiously striving fungi of the boil. The
human beings on the planet had forgotten not only the ship but very
nearly everything—how they came to this world, the use of metals,
the existence of fire, and even the fact that there was such a thing
as sunlight. They lived in the lowlands, deep under the cloud-bank,
amid surroundings which were riotous, swarming, frenzied horror.
They had become savages. They were less than savages. They had
forgotten their high destiny as men.

Dawn came. Grayness appeared overhead and increased. That was

all. The sky was a blank, colorless pall, merely mottled where the
clouds clustered a little thicker or a little thinner, as clouds do. But
the landscape was variegated enough! Where the little group of
people huddled together, there was a wide valley. Its walls rose up
and up into the very clouds. The people had never climbed those
hillsides.
They had not even traditions of what might lie above them, and
their lives had been much too occupied to allow of speculations on
cosmology. By day they were utterly absorbed in two problems
which filled every waking minute. One was the securing of food to
eat, under the conditions of the second problem, which was that of
merely staying alive.
There was only one of their number who sometimes thought of
other matters, and he did so because he had become lost from his
group of humans once, and had found his way back to it. His name
was Burl, and his becoming lost was pure fantastic accident, and his
utilization of a fully inherited power to think was the result of
extraordinary events. But he still had not the actual habit of thinking.
This morning he was like his fellows.
All of them were soaked with wetness. During the night—every night
—the sky dripped slow, spaced, solemn water-drops during the
whole of the dark hours. This was customary. But normally the
humans hid in the mushroom-forests, sheltered by the toadstools
which now grew to three man-heights. They denned in small
openings in the tangled mass of parasitic growths which flourished in
such thickets. But this last night they had camped in the open. They
had no proper habitations of their own. Caves would have been
desirable, but insects made use of caves, and the descendants of
insects introduced untold centuries before had shared in the size-
increase of paramecium and yeasts and the few true plants which
had been able to hold their own. Mining-wasps were two yards long,
and bumble-bees were nearly as huge, and there were other
armored monstrosities which also preferred caves for their own
purposes. And of course the humans could not build habitations,
because anything men built to serve the purpose of a cave would
instantly be preempted by creatures who would automatically
destroy any previous occupants.
The humans had no fixed dens at any time. Now they had not even
shelter. They lacked other things, also. They had no tools save
salvaged scraps of insect-armor—great sawtoothed mandibles or
razor-pointed leg-shells—which they used to pry apart the edible
fungi on which they lived, or to get at the morsels of meat left
behind when the brainless lords of this planet devoured each other.
They had not even any useful knowledge, except desperately
accurate special knowledge of the manners and customs of the
insects they could not defy. And on this special morning they
concluded that they were doomed. They were going to be killed.
They stood shivering in the open, waiting for it to happen.
It was not exactly news. They had had warning days ago, but they
could do nothing about it. Their home valley, to be sure, would have
made any civilized human being shudder merely to look at it, but
they had considered it almost paradise. It was many miles long, and
a fair number wide, and a stream ran down its middle. At the lower
end of the valley there was a vast swamp, from which at nightfall
the thunderously deep-bass croaking of giant frogs could be heard.
But that swamp had kept out the more terrifying creatures of that
world. The thirty-foot centipedes could not cross it or did not choose
to. The mastodon-sized tarantulas which ravaged so much of the
planet would not cross it save in pursuit of prey. So the valley was
nearly a haven of safety.
True, there was one clotho spider in its ogre's castle nearby, and
there was a labyrinth spider in a minor valley which nobody had ever
ventured into, and there were some—not many—praying-mantises
as tall as giraffes. They wandered terribly here and there. But most
members of insect life here were absorbed in their own affairs and
ignored the humans. There was an ant-city, whose foot-long warriors
competed with the humans as scavengers. There were the bees,
trying to eke out a livelihood from the great, cruciform flowers of the
giant cabbage-plants and the milkweeds when water-lilies in the
swamps did not bear their four-foot blooms. Wasps sought their own
prey. Flies were consumers of corruption, but even the flies two feet
in length would shy away from a man who waved his arms at it. So
this valley had seemed to these people to be a truly admirable place.
But a fiend had entered it. As the gray light grew stronger the
shivering folk looked terrifiedly about them. There were only twenty
of the people now. Two weeks before there had been thirty. In a
matter of days or less, there would be none. Because the valley had
been invaded by a great gray furry spider!

There was a stirring, not far from where the man-folk trembled.
Small, inquisitive antennae popped into view among a mass of large-
sized pebbles. There was a violent stirring, and gravel disappeared.
Small black things thrust upward into view and scurried anxiously
about. They returned to the spot from which they had emerged.
They were ants, opening the shaft of their city after scouting for
danger outside. They scratched and pulled and tugged at the plug of
stones. They opened the ant-city's artery of commerce. Strings of
small black things came pouring out. They averaged a foot in length,
and they marched off in groups upon their divers errands. Presently
a group of huge-jawed soldier-ants appeared, picking their way
stolidly out of the opening. They waited stupidly for the workers they
were to guard. The workers came, each carrying a faintly greenish
blob of living matter. The caravan moved off. The humans knew
exactly what it was. The green blobs were aphids—plant lice: ant-
cows—small creatures sheltered and guarded by the ants and daily
carried to nearby vegetation to feed upon its sap and yield
inestimable honeydew.
Something reared up two hundred yards away, where the thin mist
that lay everywhere just barely began to fade all colorings before it
dimmed all outlines. The object was slender. It had a curiously
humanlike head. It held out horrible sawtoothed arms in a gesture
as of benediction—which was purest mockery. Something smaller
was drawing near to it. The colossal praying mantis held its pose,
immovable. Presently it struck downward with lightning speed. There
was a cry. The mantis rose erect again, its great arms holding
something that stirred and struggled helplessly, and repented its
unconsonanted outcry. The mantis ate it daintily as it struggled and
screamed.
The humans did not watch this tragedy. The mantis would eat a
man, of course. It had. The only creatures immune to its menace
were ants, which for some reason it would not touch. But it was a
mantis' custom after spotting its prey to wait immobile for the
unlucky creature to come within its reach. It preferred to make its
captures that way. Only if a thing fled did the mantis pursue with
deadly ferocity. Even then it dined with monstrous deliberation as
this one dined now. Still, mantises could be seen from a distance
and hidden from. They were not the terror which had driven the
humans even from their hiding-places.
It had been two weeks since the giant hunting-spider had come
through a mountain pass into this valley to prey upon the life within
it. It was gigantic even of its kind. It was deadliness beyond
compare. The first human to see it froze in terror. It was disaster
itself. Its legs spanned yards. Its fangs were needle-sharp and feet
in length—and poisoned. Its eyes glittered with insatiable, insane
blood-lust. Its coming was ten times more deadly to the unarmed
folk than a Bengal tiger loose in the valley would have been.
It killed a man the very first day it was in the valley, leaving his
sucked-dry carcass, and going on to destroy a rhinoceros-beetle and
a cricket—whose deep-bass cries were horrible—and proceeded
down the valley, leaving only death behind it. It had killed other men
and women since. It had caught four children. But even that was not
the worst. It carried worse, more deadly, more inevitable disaster
with it.
Because, bumping and bouncing behind its abdomen as it moved,
fastened to its body with cables of coarse and discolored silk, the
hunting-spider dragged a burden which was its own ferocity many
times multiplied. It dragged an egg-bag. The bag was larger than its
body, four feet in diameter. The female spider would carry this
burden—cherishing it—until the eggs hatched. Then there would be
four to five hundred small monsters at large in the valley. And from
the instant of their hatching they would be just such demoniac
creatures as their parents. They would be small, to be sure. Their
legs would span no more than a foot. Their bodies would be the size
of a man's fist. But they could leap two yards, instantly they reached
the open air, and their inch-long fangs would be no less envenomed,
and their ferocity would be in madness, in insanity and in stark
maniacal horror equal the great gray fiend which had begot them.
The eggs had hatched. Today—now—this morning—they were
abroad. The little group of humans no longer hid in the mushroom-
forests because the small hunting-spiders sought frenziedly there for
things to kill. Hundreds of small lunatic demons roamed the valley.
They swarmed among the huge toadstools, killing and devouring all
living things large and small. When they encountered each other
they fought in slavering, panting fury, and the survivors of such
duels dined upon their brothers. Small truffle-beetles died, clicking
futilely. Infinitesimal grubs, newly hatched from butterfly eggs and
barely six inches long, furnished them with tidbits. But they would
kill anything and feast upon it.
A woman had died yesterday, and two small gray devils battled
murderously above her corpse.
Just before darkness a huge yellow butterfly had flung itself
agonizedly aloft, with these small dark horrors clinging to its body,
feasting upon the juices of the body their poison had not yet done to
death.
And now, at daybreak, the humans looked about despairingly for
their own deaths to come to them. They had spent the night in the
open lest they be trapped in the very forests that had been their
protection. Now they remained in clear view of the large gray
murderer should it pass that way. They did not dare to hide because
of that ogreish creature's young, who panted in their blood-lust as
they scurried here and there and everywhere.
As the day became established, the clouds were gray—gray only.
The night-mist thinned. One of the younger women of the tribe—a
girl called Saya—saw the huge thing far away. She cried out,
choking. The others saw the monster as it leaped upon and
murdered a vividly colored caterpillar on a milkweed near the limit of
vision. The milkweed was the size of a tree. The caterpillar was four
yards long. While the enormous victim writhed as it died, not one of
the humans looked away. Presently all was still. The hunting-spider
crouched over its victim in obscene absorption. Having been
madness incarnate, it now was the very exemplar of a horrid
gluttony.
Again the humans shivered. They were without shelter. They were
without even the concept of arms. But it was morning, and they
were alive, and therefore they were hungry. Their desperation was
absolute, but desperation to some degree was part of their lives. Yet
they shivered and suffered. There were edible mushrooms nearby,
but with the deadly small replicas of the hunting-spider giant
roaming everywhere, any movement was as likely to be deadly as
standing still to be found and killed. The humans murmured to one
another, fearfully.
But there was the young man called Burl, who had been lost from
his tribe and had found it again. The experience had changed him.
He had felt stirrings of atavistic impulses in recent weeks—the more
especially when the young girl Saya looked at him. It was not
normal, in humans conditioned to survive by flight, that Burl should
feel previously unimagined hunger for fury—a longing to hate and do
battle. Of course men sometimes fought for a particular woman's
favor, but not when there were deadly insects about. The
carnivorous insects were not only peril, but horror unfaceable. So
Burl's sensations were very strange. On this planet a courtship did
not usually involve displays of valor. A man who was a more skillful
forager than the foot-long ants was an acceptable husband. Warriors
did not exist.
Burl did not even know what a warrior was. Yet today the sullen,
unreasonable impulses to conduct what he could not quite imagine
were very strong. He knew all the despairing terror the others felt.
But he also was hungry. The sheer doom that was upon his group
did not change the fact that he wanted to eat, nor did it change the
fact that he felt queer when the girl Saya looked at him. Because
she was terrified, the same sort of atavistic process was at work in
her. She looked to Burl. Men no longer served as protectors against
enemies so irresistible as giant spiders. It was not possible. But
when Burl realized her regard his chest swelled. He felt a half-
formed impulse to beat upon it. His new-found reasoning processes
told him that this particular fear was different in some fashion from
the terrors men normally experienced. It was. This was a different
sort of emergency. Most dangers were sudden and either
immediately fatal or somehow avoidable. This was different. There
was time to savor its meaning and its hopelessness. It seemed as if
it should be possible to do something about it. But Burl was not
able, as yet, to think what to do. The bare idea of doing anything
was unusual, now. Because of it, though, Burl was able to disregard
his terror when Saya regarded him yearningly.

The other men muttered to each other of the sudden death in the
mushroom thickets. No less certain death now feasted on the dead
yellow caterpillar. But Burl abruptly pushed his way clear of the small
crowd and scowled for Saya to see. He moved toward the nearest
fungus-thicket. An edible mushroom grew at its very edge. He
marched toward it, swaggering. Men did not often swagger on this
planet.
But then he ceased to swagger. His approach to the mingled mass of
toadstools and lesser monstrosities grew slower. His feet dragged.
He came to a halt. His impulse to combat conflicted with the facts of
here and now. His flesh crawled at the thought of the grisly small
beasts which now might be within yards. These thickets had been
men's safest hiding-places. Now they were places of surest disaster.
He stopped, with a coldness at the pit of his stomach. But as it was
a new experience to be able to have danger come in a form which
could be foreseen, so Burl now had a new experience in that he was
ashamed to be afraid. Somehow, having tacitly undertaken to get
food for his companions, he could not bring himself to draw back
while they watched. But he did want desperately to get the food in a
hurry and get away from there.
He saw a gruesome fragment of a tragedy of days before. It was the
emptied, scraped, hollow leg-shell of a beetle. It was horrendously
barbed. Great, knife-edged spines lined its edge. They were six
inches in length. And men did not have weapons any more, but they
sometimes used just such objects as this to dismember defenseless
giant slugs they came upon.
Burl picked up the hollow shell of the leg-joint. He shook it free of
clinging moulds—and small things an inch or two in length dropped
from it and scurried frantically into hiding. He moved hesitantly
toward the edible mushroom which would be food for Saya and the
rest. He was four yards from the thicket. Three. Two. He needed to
move only six feet, and then slice at the flabby mushroom-head, and
he would be at least an admirable person in the eyes of Saya.
Then he cried out thinly. Something small, with insane eyes, leaped
upon him from the edge of a giant toadstool.
It was, of course, one of the small beasts which had hatched from
the hunting-spider's egg-bag. It had grown. Its legs now spanned
sixteen inches. Its body was as large as Burl's two fists together. It
was big enough to enclose his head in a cage of loathesomeness
formed by its legs, while its fangs tore at his scalp. Or it could cover
his chest with its abominableness while its poison filled his veins,
and while it feasted upon him afterward....
He flung up his hands in a paralytic, horror-stricken attempt to ward
it off. But they were clenched. His right hand did not let go of the
leg-section with its razor-sharp barbs.
The spider struck the beetle-leg. He felt the impact. Then he heard
gaspings and bubblings of fury. He heard an indescribable cry which
was madness itself. The chitinous object he had picked up now
shook and quivered of itself.
The spider was impaled. Two of its legs were severed and twitched
upon the ground before him. Its body was slashed nearly in half. It
writhed and struggled and made beastly sounds. Thin, colored fluids
dripped from it. A disgusting musky smell filled the air. It strove to
reach and kill him as it died. Its eyes looked like flames.
Burl's arm shook convulsively. The small thing dropped to the
ground. Its remaining legs moved frantically but without purpose.
It died, though its leg continued to twitch and stir and quiver.
Burl remained frozen, for seconds. It was an acquired instinct; a
conditioned reflex which humans had to develop on this world.
When danger was past, one stayed desperately still lest it return. But
Burl's thoughts were now not of horror but a vast astonishment. He
had killed a spider! He had killed a thing which would have killed
him! He was still alive!
And then, being a savage, and an animal, as well as a human being,
he acted according to that highly complicated nature. As a savage,
he knew with strict practicality that it was improbable that there was
another baby spider nearby. If there had been, they would have
fought each other. As an animal, he was again hungry. As a human
being, he was vain.
So he moved closer to the toadstool-thicket and put his hand out
and broke off a great mass of the one edible mushroom at the edge.
A noisesome broth poured out and little maggots dropped to the
ground and writhed there in it. But most of what he had broken off
was sound. He turned to take it to Saya. Then he saw the dropped
weapon and the spider. He picked up the weapon.
The spider's legs still twitched, though futilely. He spiked the small
body on the beetle-leg's spines. He strode back to the remnant of
his tribe with a peculiar gait that even he had not often practiced.
It was rather more pronounced than a swagger. It was a strut.
They trembled when they saw the dead creature he had killed. He
gave Saya the food. She took it, looking at him with bright and
intense eyes. He took a part of the mushroom for himself and ate it,
scowling. Thoughts were struggling to form in his mind. He was not
accustomed to thinking, but he had done more of it than any other
of the pitiful group about him.
He felt eyes watching him. There were five adult men in this group
besides himself, and six women. The rest were children, from
gangling adolescents to one mere infant in arms. They were a
remarkably colorful group at the moment, had he only known it. The
men wore yellow-and-gold-brown loin-cloths of caterpillar-fur,
stripped from the drained carcasses of creatures that the formerly
resident clothed spider had killed. The women wore cloaks of
butterfly-wing, similarly salvaged from the remnants of a meal left
unfinished by a finicky or engorged praying mantis. The stuff was
thick and leathery, but it was magnificently tinted in purples and
yellows.
Time passed. The mushroom Burl had brought was finished. Some
eyes always explored the clear ground around this group. But other
eyes fixed themselves upon Burl. It was not a consciously
questioning gaze. It was surely not a hopeful one. But men and
women and children looked at him. They marveled at him. He had
dared to go and get food! He had been attacked by one of the
creatures who doomed them all, but he was not dead! Instead, he
had killed the spider! It was marvelous! It was unparalleled that a
man should kill anything that attacked him!

The doomed small group regarded Burl with wondering eyes. He

brushed his hands together. He looked at Saya. He wished to be
alone with her. He wished to know what she thought when she
looked at him. Why she looked at him. What she felt when she
looked at him.
He stood up and said dourly:
"Come!"
She moved timidly and gave him her hand. He moved away. There
was but one way that any human being on this planet would think to
move, from this particular spot just now—away from the still-
feasting gigantic horror whose offspring he had killed. The folk
shivered near the edge of the first upward slope of the valley wall.
Burl moved in that direction. Toward the slope. Saya went with him.
Before they had gone ten yards a man spoke to his wife. They
followed Burl, with their three children. Five yards more, and two of
the remaining three adult men were hustling their families in his
wake also. In seconds the last was in motion.
Burl moved on, unconscious of any who followed him, aware only of
Saya. The procession, absurd as it was, continued in his wake simply
because it had begun to do so. A skinny, half-grown boy regarded
Burl's stained weapon. He saw something half-buried in the soil and
moved aside to tug at it. It was part of the armor of a former
rhinoceros-beetle. He went on, rather awkwardly holding a weapon
which might have been called a dagger, eighteen inches long, except
that no dagger would have a hand-guard nearly its own length in
diameter.
They passed a struggling milkweed plant, no more than twenty feet
high and already scabrous with scale and rusts upon its lower parts.
Ants marched up and down its stalk in a steady, single file, placing
aphids from the ant-city on suitable spots to feed, and to multiply as
only parthenogenic aphids can do. But already on the far side of the
milkweed, an ant-lion climbed up to do murder among them. The
ant-lion was the larval form the lace-wing fly, of course. Aphids were
its predestined prey.
Burl continued to march, holding Saya's hand. The reek of formic
acid came to his nostrils. But that was only ants. The slope grew
steeper. Massacre began behind him on the tree-sized milkweed.
The ant-lion which even when it was but half an inch long, on Earth,
could bite through the skin of a man—the ant-lion reached the
pasturing cows. It plunged into slaughter. It was demoniac. It was
such ghastly ferocity that the eggs from which its kind hatched were
equipped, each one, with a plastic column to hold it well away from
the object on which the clutch of eggs were laid. But for this
precaution by the maternal lace-wing fly, the first of her brood to
hatch would devour its unhatched brothers and sisters. This ant-lion
charged into the placidly feeding aphids on the milkweed plant. It
seized one and crushed it, holding it aloft so that the juices of its
body would pour into the ant-lion's mouth. Almost instantly, it
seemed, the mild-eyed aphid was a shrunken empty sack. The ant-
lion seized another. The remaining aphids fed placidly while their
enemy did vast slaughter among them.
Clickings and a shrill stridulation sounded. Warrior-ants climbed with
stupid ferocity to offer battle.
Burl moved on to a minor eminence. He reached its top and looked
sharply about him with the caution that was the price of existence
on this world. Two hundred feet away, a small scurrying horror raged
and searched among the rough-edged layers of what on other
worlds was called paper-mould or rock-tripe. Here it was thick as
quilting, and infinitesimal creatures denned under it. The sixteen-
inch spider devoured them, making gluttonous sounds. But it was
busy, and all spiders are relatively short-sighted.
Burl turned to Saya—and realized that all the human folk had
followed him. One of the adults was reaching fearfully for part of a
discarded cricket-shell in the ground. He tore free an emptied, sickle-
shaped jaw. It was curved and sharp and deadly if properly wielded.
The man had seen Burl kill something. He tried vaguely to imagine
killing something himself. He was not too successful. Another man
tugged at the ground. The skinny boy was practicing thrusts with his
giant dagger.
Two of the adults were armed, without any clear idea of what to do
with their arms. But Burl knew, now.
He regarded them angrily. He had not meant to desert them, or
even to take Saya permanently from among them. Humans had little
enough of satisfaction on this planet. The scared company of their
kind was one of the most important. So Burl did not resent that they
had followed him. He did resent that they were near when he
wanted to talk to Saya in what he did not yet think of as lover-like
seclusion.
They halted, regarding him humbly. They had been hungry, and he
had found food for them. They had been paralyzed by terror, and he
had dared to move. So they moved with him. They might have
followed anybody else, but only Burl had initiative—so far. They
trustfully waited to follow and to imitate him for so long as panic
numbed their ability to think for themselves.
Burl opened his mouth to shout furiously at them. But it was not a
good idea for humans to draw attention. Spiders did not hunt by
scent, but sound sometimes drew them. Burl closed his mouth
again, in a taut straight line. The men looked at him supplicatingly.
They had never been lost, and so had never learned to think even a
little. Burl had learned to think in a rudimentary fashion and now he
suddenly perceived that it was pleasing to have all the tribe regard
him so worshipfully, even if not in quite the same fashion as Saya.
He was suddenly aware that even as Saya had obeyed him when he
told her to come with him, they would obey. He had, at the moment,
no commands to give, but he immediately invented one for the
pleasure of seeing it carried out.
"I carry sharp things," he said sternly. "I killed a spider. Go find
sharp things to carry."
They were a meek and abject folk, and they were desperately in
need of something to do to take their minds from the uselessness of
doing anything at all.
They moved to obey. Saya would have loosened her hand and
obeyed, too, but Burl held her beside him. One of the women, with a
child three years old, laid the child down by Burl's feet while she
went fearfully to seek some fragment of a dead creature, that would
meet Burl's specification of sharpness.
Burl heard a stifled scream. A ten-year-old boy stood paralyzed,
staring in an agony of horror at something which had stepped from
behind a misshapen fungoid object.
It was a pallidly greenish creature with a small head and enormous
eyes. It was a very few inches taller than a man. Its abdomen
swelled gracefully into a pleasing, leaf-like shape. The boy faced it,
paralyzed by horror, and it stood stock-still. Its great, hideously spiny
arms were spread out in a pose of pious benediction.
 "The boy faced it, paralyzed by horror."

It was a partly-grown praying mantis, not very long hatched. It

stood rigid, waiting benignly for the boy to come closer. If he fled, it
would fling itself after him with ferocity beside which the fury of a
tiger would seem kittenish. If he approached, its fanged arms would
flash down, pierce his body, and hold him inextricably fast by the
spikes that were worse than trap-claws. And of course it would not
wait for him to die before it began its meal.
The small party of humans stood frozen. They were filled with horror
for the boy. They were cast into a deep abyss of despair by the sight
of a half-grown mantis, because if there was one such miniature
insect-dinosaur in the valley, there would be many others. Hundreds
of others. This meant there had been a hatching of them. And they
were as deadly as spiders.

But Burl did not think in such terms just now. Vanity filled him. He
had commanded, and he had been obeyed. But now obedience was
forgotten because there was this young praying mantis. If men had
ever thought of fighting such a creature, it could have destroyed any
number of them by pure ferocity and superiority of armament. But
Burl raged. He ran toward the spot. Even mantises were sometimes
frightened by the unexpected. Burl seized a lumpish object barely
protruding from the ground. It looked like a rock. It was actually a
flattened ball-fungus, feeding on the soil through thin white threads
beneath it. Burl wrenched it free and hurled it furiously at the young
monster.
Insects simply do not think. Something came swiftly at it, and the
mantis flashed its ghastly arms to seize and kill its attacker. The ball-
fungus was heavy. It literally knocked the mantis backward. The boy
fled frantically. The insect fought crazily against the thing it thought
had assailed it.
The humans gathered around Burl hundreds of yards away—again
uphill. The slope of the mountain-flank was marked here. They
gathered about Burl because of an example set by the woman who
had left her three-year-old child behind. Saya, in the unfailing
instinct of a girl for a small child, had snatched it up when Burl left
her. Then she had joined him because the instinct which had made
her obey him in starting off—it was not quite the same instinct which
moved the others—also bade her follow him wherever he went. The
mother of the child went to retrieve her deposit. Other figures
moved cautiously toward him. The tribe was reconvened.
The floor of the valley seemed a trifle obscured. The mist that hung
always in the air made it seem less distinct; less actual; not quite as
real as it had been.
Burl gulped and said sternly:
"Where are the sharp things?"
The men looked at one another, numbly. Then one spoke
despairingly, ignoring Burl's question. "Now," said the man dully,
"there was not only the hunting-spider in the valley, but its young.
And not only the young of the hunting-spider, but the young of a
mantis ... It was hard to stay alive at the best of times. Now it had
become impossible ..."
Burl glared at him. It was neither courage nor resolution. He had
come to realize what a splendid sensation it was to be admired by
one's fellows. The more he was admired, the better. He was enraged
that people thought to despair.
"I," said Burl haughtily, "am not going to stay here. I go to a place
where there are neither spiders nor mantises. Come!"
He held out his hand to Saya. She gave the child to its mother and
look his hand. Burl stalked haughtily away, and she went with him.
He went uphill. Naturally. He knew there were spiders and mantises
in the valley. So many that to stay there was to die. So he went
away from where they were.
Burl had found out that adulation was enjoyable and authority
delectable. He had found that it was pleasant to be a dictator. And
then he had been disregarded. So he marched furiously away from
his folk, in exactly the fashion of a spoiled child refusing to play any
longer. He happened to march up the mountainside toward the
cloud-bank that he considered the sky. He had no conscious intent
to climb the mountain. He did not intend to lead the others. He
meant to sulk, by punishing them through the removal of his own
admirable person from their society. But they followed him.
So he led his people upward. It has happened on other planets, in
other manners. Most human achievements come about through the
daring of those who strive.

The sun was very near. It shone upon the top of the cloud-bank and
the clouds glowed with a marvelous whiteness. It shone upon the
mountain-peaks where they penetrated the clouds, and the peaks
were warmed, and there was no snow anywhere despite the height.
There were winds here where the sun shone. The sky was very blue.
At the edge of the plateau where the cloud-bank lay below, the
mountainsides seemed to descend into a sea of milk. Great
undulations in the mist had the seeming of waves, which moved
with great deliberation toward the shores. They seemed sometimes
to break against the mountain-wall where it was cliff-like, and
sometimes they seemed to flow up gentler inclinations like water
flowing up a beach.
All this was in the slowest of slow motion, because the cloud-waves
were sometimes miles from crest to crest.
The look of things was different on the plateau, too. This part of the
unnamed world, no less than the lowlands, had been seeded with
life on two separate occasions. Once with bacteria and moulds and
lichens to break up the rocks and make soil of them, and once with
seeds and insects-eggs and such living things as might sustain
themselves immediately upon hatching. But here on the heights the
conditions were drastically unlike the lowland tropic moisture.
Different things had thriven, and in quite different fashion.
Here moulds and yeasts and rusts were stunted by the sunlight.
Grasses and weeds and trees survived, instead. This was an ideal
environment for plants that needed sunlight to form chlorophyll, and
chlorophyll to make use of the soil that had been formed. So here
was vegetation that was nearly Earth-like. And there was a
remarkable side-effect on the fauna which had been introduced at
the same time and in the same manner as down below. In coolness
which amounted to a temperate climate there could be no such
frenzy of life as formed the nightmare-jungles in the lowlands. Plants
grew at a slower tempo than fungi, and less luxuriantly. There was
no adequate food-supply for large-sized plant-eaters. Insects which
were to survive in sunshine could not grow to be monsters.
Moreover, the nights were chill. Many insects grow torpid in the cool
of a temperate-zone night, but warm up to activity soon after
sunrise. But a large creature, made torpid by cold, will not revive so
quickly. If large enough, it will not become fully active until close to
dusk. On the plateau, the lowland monsters would starve in any
case. But more—they would have only a fraction of a day of full
activity.
There was a necessary limit then, to the size of the insects that lived
above the clouds. The life on the plateau would not have seemed
horrifying at all to humans living on other planets. Save for the
absence of birds to sing and lack of a variety of small mammals, the
untouched sunlit plateau with its warm days and briskly chill nights
would have impressed most men as an ideal habitation.
But Burl and his companions were hardly prepared to see it that way
at first glimpse. Certainly if told about it beforehand, they would
have viewed it with despair.
But they did not know beforehand. They toiled upward, their leader
moved by such ridiculous motives as have sometimes caused men to
achieve greatness throughout all history. Back on Earth, two great
continents were discovered by a man trying to get spices to conceal
the gamey flavor of half-spoiled meat. The power that drives mile-
long space-craft, and that lights and runs the cities of the galaxy,
was first developed because it could be used in bombs to kill other
men. There were precedents for Burl leading his fellows into
sunshine merely because he was angered that they ceased to
admire him.
The trudging, climbing folk were high above the valley, now. The
thin mist that was never absent anywhere had hidden their former
home, little by little. They climbed a steeply slanting mountain-flank.
The stone was mostly covered by ragged, bluish-green rock-tripe in
partly overlapping sheets. Such stuff is always close behind the
bacteria which first attack a rock-face. On a slope, it clings while soil
is washed downward as fast as it forms. The people never ate it. It
produced frightening cramps. In time they would learn that if
thoroughly dried it can he soaked to pliability again and cooked to a
reasonable palatability. But so far they knew neither dryness nor fire.
Nor had they ever known such surroundings as presently enveloped
them. A slanting, stony mountainside which stretched up
frighteningly to the very sky. Grayness overhead. Grayness, also, to
one side—the side away from the mountain. And equal grayness
below. The valley in which they lived could no longer be seen at all.
Trudging and scrambling up the interminable incline, the people of
Burl's personal following gradually realized the strangeness of their
surroundings. As one result, they grew sick and dizzy. To them it
seemed that the solid earth had tilted, and might presently tilt
further. There was no horizon, but they had never seen a horizon. So
they felt that what had been down was now partly behind, and they
feared lest a turning universe let them fall ultimately toward the
grayness they considered sky.
In this frightening strangeness, their only consolation was the
company of their fellows. To stop would be to be abandoned in this
place where all values were turned topsy-turvy. To go back—but
none of them could imagine descending again to be devoured as
one-third of their number already had been. If Burl had stopped, his
followers would have squatted down and shivered together
miserably, and waited for death. They had no thought of adventure
nor any hope of safety. The only goodnesses they could imagine
were food and the nearness of other humans. They clung together,
obsessed by the dread of being left alone.
Burl's motivation was no longer noble. He had started uphill in a fit
of sulks, and he was ashamed to stop.
They came to a place where the mountain-flank sank inward. There
was a flat area, and behind it there was a winding cañon of sorts,
like a vast crack in the mountain's substance. Burl breasted the
curving edge, and walked on level ground. Then he stopped short.
The mouth of the cañon was perhaps fifty yards from the lip of the
downward slope. There was this level space, and on it there were
toadstools and milkweed, and there was food. It was a small,
isolated asylum for life such as they were used to. It could have
been that here they could have found safety. But it wasn't that way.

They saw the web at once. It was slung from between the opposite
cliff-walls by cables two hundred feet long. Its radiating cables
reached down to anchorages on stone. The snare-threads, winding
out and out in that logarithmic spiral which men on other planets
had noted thousands of years before—the snare-threads were a yard
apart. The web was set for giant game. It was empty now, but Burl
searched keenly and saw the tight-rope-cable leading from the very
center of the web to a rocky shelf some fifty feet above the cañon's
floor. At its end he saw the spider. It waited there, almost invisible
against the stone, with one furry leg touching the cable that led to
its waiting-place so that the slightest touch on any part of the web
would warn it instantly.
Burl's followers accumulated behind him. They stared. They knew, of
course, that a web-spider will not leave its snare under any normal
circumstances. They were not afraid of that. But they looked at the
ground between the web and themselves.
It was a charnel-house of murdered creatures. Half-inch-thick wing-
cases of dead beetles. The cleaned-out carcasses of other giants.
The ovipositor of an ichneumon-fly—six feet of slender, springy,
deadly-pointed tube—and abdomen-plates of bees and draggled
antennae of moths and butterflies.
Something very terrible lived in this small place. The mountainsides
were barren of food for big flying things. Anything which did fly so
high for any reason would never land on sloping, foodless stone. It
would land here. And very obviously it would die. Because
something—something—killed them as they came. It denned back in
the cañon where they could not see. It dined here.
The humans looked and shivered. All but Burl. He deliberately chose
for himself a magnificent lance grown by one dead creature for its
own defense. He pulled it out of the ground and cleaned it with his
hands. He seemed absorbed, but he was terribly aware of the inner
depths of the cañon. He was actually pretending, for the sake of
what he believed his dignity.
Fearfully, the other humans imitated him in choosing weapons from
the armory of the devoured. Then Burl stalked grandly to one side
and began to climb again. His people followed him in numbed
silence. They were filled with dread, but it was not quite terror.
Insects do not stalk their prey. The deadly unseen monster of the
cañon had not attacked them. Therefore, it did not know they were
there. And therefore they were safe from it until it appeared. But
none of them desired to stay.
The slope lessened here, and half a mile further on there was a
small thicket of mushrooms. From within it came the cheerful loud
clicking of some small beetle, arrived at this spot nobody could
possibly know how, but happily ensconced in a twenty-yard patch of
jungle above a hollow that had gathered soil through the centuries.
There were edible mushrooms in the thicket.
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