Visit https://ebookultra.
com to download the full version and
explore more ebooks or textbooks
Enzymatic Reaction Mechanisms 1st Edition Perry A.
Frey
_____ Click the link below to download _____
https://ebookultra.com/download/enzymatic-reaction-
mechanisms-1st-edition-perry-a-frey/
Explore and download more ebooks or textbooks at ebookultra.com
Here are some recommended products that we believe you will be
interested in. You can click the link to download.
Organic Reaction Mechanisms 2012 1st Edition A. C. Knipe
(Ed.)
https://ebookultra.com/download/organic-reaction-mechanisms-2012-1st-
edition-a-c-knipe-ed/
Organic Reaction Mechanisms 2014 An annual survey covering
the literature dated January to December 2014 1st Edition
A. C. Knipe
https://ebookultra.com/download/organic-reaction-mechanisms-2014-an-
annual-survey-covering-the-literature-dated-january-to-
december-2014-1st-edition-a-c-knipe/
Productive Group Work Nancy Frey
https://ebookultra.com/download/productive-group-work-nancy-frey/
Writing As a Learning Activity 1st Edition Perry Klein
https://ebookultra.com/download/writing-as-a-learning-activity-1st-
edition-perry-klein/
Essentials of Chemical Reaction Engineering 1st Edition
Fogler
https://ebookultra.com/download/essentials-of-chemical-reaction-
engineering-1st-edition-fogler/
Alzheimers Disease Advances for a New Century 1st Edition
George Perry
https://ebookultra.com/download/alzheimers-disease-advances-for-a-new-
century-1st-edition-george-perry/
SPSS explained 1st Edition Perry Hinton
https://ebookultra.com/download/spss-explained-1st-edition-perry-
hinton/
Chemical Reaction Engineering 2nd Edition Martin Schmal
(Editor)
https://ebookultra.com/download/chemical-reaction-engineering-2nd-
edition-martin-schmal-editor/
Green Reaction Media in Organic Synthesis 1st Edition
Mikami Koichi
https://ebookultra.com/download/green-reaction-media-in-organic-
synthesis-1st-edition-mikami-koichi/
Enzymatic Reaction Mechanisms 1st Edition Perry A.
Frey Digital Instant Download
Author(s): Perry A. Frey, Adrian D. Hegeman
ISBN(s): 9780195122589, 0195122585
Edition: 1
File Details: PDF, 32.44 MB
Year: 2007
Language: english
Enzymatic Reaction
Mechanisms
This page intentionally left blank
ENZYMATIC REACTION
MECHANISMS
Perry A. Frey and
Adrian D. Hegeman
1
2007
1
Oxford University Press, Inc., publishes works that further
Oxford University’s objective of excellence
in research, scholarship, and education
Oxford New York
Auckland Cape Town Dar es Salaam Hong Kong Karachi
Kuala Lumpur Madrid Melbourne Mexico City Nairobi
New Delhi Shanghai Taipei Toronto
With offices in
Argentina Austria Brazil Chile Czech Republic France Greece
Guatemala Hungary Italy Japan Poland Portugal Singapore
South Korea Switzerland Thailand Turkey Ukraine Vietnam
Copyright © 2007 by Oxford University Press
Published by Oxford University Press, Inc.
198 Madison Avenue, New York, New York, 10016
www.oup.com
Oxford is a registered trademark of Oxford University Press
All rights reserved. No part of this publication may be reproduced,
stored in a retreival system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording, or otherwise, without
the prior permission of Oxford University Press.
9 8 7 6 5 4 3 2 1
Printed in the United States of America
on acid-free paper
To Professor Frank H. Westheimer,
a great scientist, a great scholar, and a great teacher.
This page intentionally left blank
Preface
In planning this book, we gave considerable thought to how we should proceed.
Enzymology is a very large and multifaceted field that interfaces with and stimulates
research in peripheral fields such as protein structure, spectroscopy, x-ray crystallography,
polymer science, biotechnology, protein folding, cellular metabolism, and biological regula-
tion. All facets cannot be covered in a single volume. Our motivation to write about the
chemical mechanisms of enzymatic catalysis guided us in defining the scope of this
volume. To bring the material between two covers, it would be necessary to focus on the
chemical mechanisms. In making this decision, we did not intend to de-emphasize related
areas of enzymology; we simply could not give them as much attention as the reaction
mechanisms in the space available for a single volume. Many of these peripheral areas are
brought into the discussion of individual enzymes, with less detail, to place the chemical
mechanisms in biological or chemical context. However, the emphasis remains on the
mechanisms of enzymatic catalysis.
In the first five chapters, we define the scope of the problem of understanding enzy-
matic catalysis and introduce most of the principles, theory, and concepts. We begin with
principles and theories of catalysis and the role of the active site in chapter 1. We continue
with an overview of enzyme kinetics in chapter 2. In chapter 3, we discuss classic organic
coenzymes and, in chapter 4, metallocoenzymes and cofactors and their contributions to
catalyzing enzymatic processes. In chapter 5, we discuss the theory and practice of the
development and characterization of specific inhibitors and inactivators of enzymes, an
important focus in pharmaceutical and agricultural chemistry. In this part of the book, we
include a few brief case studies or sketches of the mechanisms of action of key enzymes,
including structures and chemical and spectroscopic results, to exemplify principles.
In the second part of this volume, we focus on types of enzymatic reactions in succeed-
ing chapters. Each chapter includes a discussion of the underlying chemistry and brief case
studies exemplifying enzymes in that class, including the most relevant chemical, kinetic,
and structural results pertaining to the mechanism. We start with the kinetically simplest
reactions and move toward increasing kinetic complexity. We begin with one-substrate,
irreversible reactions in chapter 6, the proteases and esterases, and continue with one-
substrate, reversible isomerases and mutases in chapter 7. In chapter 8, we discuss decar-
boxylases and carboxylases, and in chapter 9, we address the lyases in addition and elimination
viii Preface
reactions. We continue with the kinetically more complex group transfer enzymes, the
phosphotransferases and nucleotidyltransferases, in chapter 10, the ATP-dependent
synthetases in chapter 11, and the glycosyltransferases and glycosidases in chapter 12. In
chapter 13, we continue with nitrogen and sulfur transferases; in chapter 14, with carbon-
carbon ligases and cleavage enzymes; and in chapter 15, with alkyltransferases. The
oxidoreductases are the subjects of chapter 16. In chapter 17, we discuss reactions of O2
and the mechanisms of action of the oxidases and oxygenases. In chapter 18, entitled
complex Enzymes, we conclude with discussions of the most complex enzymes, the
multienzyme complexes, the modular enzymes and multienzymes proteins, and enzymes
that function in energy transduction.
The chapters were written between 2002 and 2005. We found that each chapter became
outdated within a few weeks of being written. We chose to proceed with publication because
we prefer to have a book in print than one perpetually in preparation. In a work such as this,
there are bound to be oversights and mistakes, and we take full responsibility for them. We
are receptive to readers’ comments and suggestions for improvements, including updating.
If the book has a second printing, we will attempt to incorporate corrections and suggestions.
We include brief case studies of the mechanisms of action of more than 100 enzymes
in this volume. It has not been possible to provide complete referencing of all the significant
research on all of these enzymes. We have included leading references to recent work and
selected references to early work that stimulated research on a given enzyme. In selecting
enzymes for inclusion, we were guided by the need to incorporate at least one example of
an enzyme catalyzing each reaction type. We were also influenced by our own interests,
and we seek our readers’ indulgence on this matter.
We are grateful to colleagues who contributed in various ways to this book. We are
particularly indebted to Professors W. Wallace Cleland, George H. Reed, and Brian G. Fox,
who contributed significantly to the various chapters and who read and commented on selected
chapters. They bear no responsibility for our mistakes, and we thank them for suggesting
improvements and correcting many of our errors.
Internet Enzymology Resources
Today, it would be unnecessarily limiting to work in the field of enzymology without using
internet-based resources. Because of their rapid development and their relative imperma-
nence compared with archived literature, these resources are touched on briefly and
considered in the context of the main text.
Throughout the text, database accession information is provided in two forms. First, in
each figure for which three-dimensional structural information was used, a four-character
PDB code is listed in the caption (e.g., IMGO refers to the structure coordinate file for
horse liver alcohol dehydrogenase complexed with 2,3-diflurobenzyl alcohol and NAD+).
These codes may be used to access archived structure coordinate files and header informa-
tion from the Brookhaven Protein Data Bank (PDB; Berman et al, 2000), which is hosted
by a number of organizations, including the Research Collaboratory for Structural
Bioinformatics (RCSB; http://www.rcsb.org/pdb/) and the Enzyme Structures Database
(at http://www.ebi.ac.uk/thornton-srv/database/enzymes/). Second, as each enzyme activity
in the text is introduced, the enzyme classification (EC) number is provided. The EC
number format is EC #.#.#.#, where each refers to a unique aspect of catalysis in four hier-
archic categories so that all four numbers refer to a unique activity (e.g., EC 1.1.1.1 refers
to the activity of alcohol dehydrogenase). This systematization helps to eliminate confu-
sion resulting from conflicting naming conventions, and it serves as an accession number
for enzymes database. BRENDA (http://www.brenda.uni-koeln.de), hosted by the
University of Cologne, contains a large amount of information on enzyme substrates, prod-
ucts, and inhibitors and other functional data, which may be accessed using EC numbers
or by using cross-listed preferred or alternate nomenclature (Schomburg et al, 2004). Other
searchable databases vary over time, but they tend to be maintained and updated (or at least
linked to) by several key organizations, including the National Center for Biotechnology
Information (NCBI) (http://www.ncbi.nim.nih.gov) and the European Molecular Biology
Laboratory–European Bioinformatics Institute (EMBL-EBI) http://www.ebi.ac.uk/). These
sites frequently include access to free, often experimental, and downloadable or web-based
tools.
The PDB access sites also provide freeware for structure file viewing, although struc-
ture viewers are generally widely available. All of the structural figures presented in the
text were generated using the free visualization and rendering package MolView (vl.5.0)
x Internet Enzymology Resources
(Smith, 1995). An extensive collection of other web-based tools for protein structural and
proteomic analysis is available at the ExPASy (Expert Protein Analysis System)
proteomics server of the Swiss Institute of Bioinformatics (SIB). Many interesting exper-
imental tools, such as the web-based homology modeling package SWISS-MODEL
(http://swissmodel.expasy.org/) exist that may be of interest to anyone exploring the vast
reams of genomic data that have been accumulated in the past decade.
Finally, some novel strategies for populating and querying database have been developed
that apply specifically to the interface of high-throughput structural biology and enzymology.
The catalytic Site Atlas (CSA; http://www.ebi.uk/thornton-srv/database/CSA), for example,
provides catalytic residue annotation for enzymes in the PDB, with the ultimate goal being
the characterization of enzyme structural elements (e.g., active-site residues) that directly
pertain to function (Porter et al, 2004).
If successful, this sort of analysis may provide additional insight into the function of
uncharacterized gene products that cannot be gained from the analysis of amino acid
sequence or domain structure alone.
References
Berman HM, J Westbrook, Z Feng, G Gilliland, TN Bhat, H Weissig, IN Shindyalov, and PE
Bourne (2000) Nucleic Acids Res 28, 235.
Gasteiger E, A Gattiker, C Hoogland, I Ivanyi, RD Appel, and A Bairoch (2003) Nucleic Acids
Res 31, 3784.
Porter CT, GJ Bartlett, and JM Thornton (2004) Nucleic Acids Res 32, D129.
Schomburg I, A Chang, C Ebeling, M Gremse, C Heldt, G Huhn, and D Schomburg (2004)
Nucleic Acids Res 32, D431.
Smith TJ (1995) J Mol Graphic 13, 122.
Contents
Chapter 1. Enzymes and Catalytic Mechanisms, 1
Catalysis and the Active Site, 1
Rate Enhancement in Enzymatic Catalysis, 3
Conformational Mobility in Catalysis, 5
Substrate-Induced Conformational Changes, 5
Catalysis of Multistep Reactions, 6
Structural Mobility in Enzymes, 6
Acid-Base Catalysis, 9
Acids and Bases, 9
Acid- and Base-Catalyzed Reactions, 11
Nucleophilic Catalysis, 16
Electrophilic Catalysis, 21
Catalysis of Enolization, 21
Imine Formation by Lysine, 23
Catalysis by Metal Ions, 26
Hydrogen Bonding, 30
Strong and Weak Hydrogen Bonds, 30
Hydrogen Bonding in Catalysis, 32
Binding Energy in Catalysis, 34
Binding and Activation Energy, 34
The Active Site as an Entropy Trap, 36
Dissecting the Binding Effect in Enzymatic Action, 40
Stabilization of the Transition State, 41
Binding the Near Attack Conformation, 46
Destabilization of Ground States, 48
Rate Enhancement through Binding of Remote Groups, 48
Characterization of Active Sites, 53
xii Contents
Competitive Inhibitors: Analogs of Substrates, 53
Group-Selective Chemical Modification, 53
Site-Directed Mutagenesis, 57
Affinity Labeling, 59
Why Are Enzymes Large Molecules?, 62
Sizes of Enzymatic Binding Domains, 62
Catalytic Antibodies, 63
Chapter 2. Kinetics of Enzymatic Reactions, 69
Steady-State Kinetics, 69
One-Substrate Reactions, 70
Two-Substrate Reactions, 74
Three-Substrate Reactions, 89
Isotope Effects, 91
Classes of Isotope Effects, 91
Measurement of Isotope Effects, 95
Transient-Phase Kinetics, 101
Reaction Characteristics, 101
Transient Methods, 102
pH-Rate Profiles, 111
Profile Interpretation, 111
Measurements of pH-Rate Profiles, 111
Allosteric Regulation, 117
Theory, 118
Binding Equations for Cooperative Systems, 120
Aspartate Transcarbamoylase, 123
Chapter 3. Coenzymes I: Organic Coenzymes, 129
Nicotinamide Coenzymes, 129
Structures and Functions of Nicotinamide Coenzymes, 129
Stereospecificity of Hydride Transfer, 132
NAD+ as a Coenzyme, 134
Thiamine Pyrophosphate, 141
Structure, 141
Reaction Mechanism, 141
α-Lipoamide, 147
Pyridoxal-5′-Phosphate, 148
Enzymatic Reactions Facilitated by Pyridoxal-5′-Phosphate, 149
Pyridoxal-5′-Phosphate–Stabilized Amino Acid Carbanions, 149
Mechanisms of Pyridoxal-5′-Phosphate–Dependent Reactions, 151
Flavin Coenzymes, 158
Structures of Flavin Coenzymes, 158
Mechanisms of Flavin Catalysis, 159
Biotin, 163
Structure and Role as a Carboxyl Carrier, 163
Contents xiii
Chemistry of Biotin and N1-Carboxybiotin, 164
Mechanism of Biotin-Dependent Carboxylation, 164
Phosphopantetheine Coenzymes, 165
Structures of Phosphopantetheine Coenzymes, 165
Mechanism of Phosphopantetheine Action, 165
Folate Compounds, 167
Folate Compounds of One-Carbon Metabolism, 168
Enzymes in Tetrahydrofolate Metabolism, 170
Biological Importance of Folate, 171
Amino Acid–Based Coenzymes, 172
Pyruvoyl Decarboxylases, 172
Methylidene Imidazolinone–Dependent Deaminases, 173
Quinoproteins, 174
Chapter 4. Coenzymes II: Metallic Coenzymes, 189
Vitamin B12 Coenzymes, 190
Chemistry of B12 Coenzymes, 190
Adenosylcobalamin-Dependent Enzymes, 193
Methylcobalamin-Dependent Enzymes, 199
Heme Coenzymes, 201
Chemistry of Oxygen and Heme, 201
Heme Enzymes, 204
Oxygen Binding and Electron Transfer, 209
Mononuclear Nonheme Iron, 210
Monooxygenases, 210
Dioxygenases, 217
Oxo-Fe2 Complexes, 217
Structures, 218
Reactions of Di-iron Enzymes, 219
Metallopterin Enzymes, 222
Molybdopterin and Tungstopterin, 222
Iron-Sulfur Centers, 227
Structures, 227
Catalytic Functions, 230
S-Adenosylmethionine and Iron-Sulfur Centers, 234
Catalytic Action of S-Adenosylmethionine and
[4Fe–4S] Centers, 234
Stoichiometric Reactions of S-Adenosylmethionine
and [4Fe–4S] Centers, 236
Divalent Metal Ions, 237
Electrostatic Activation of Coordinated Water, 237
Electrostatic Activation of Enolization, 238
Copper as a Cofactor, 240
Copper Proteins, 240
xiv Contents
Other Copper Enzymes, 241
Nickel Coenzymes, 243
Nickel in Methanogenesis, 243
Other Nickel Coenzymes, 245
Long-Range Electron Transfer, 247
Biological Electron Transfer, 247
Marcus Theory, 248
Chapter 5. Enzyme Inhibition, 253
Two-Substrate Analogs, 254
Inhibition and Binding, 254
PALA and Aspartate Transcarbamylase, 254
Suicide Inactivation, 255
Thymidylate Synthase, 255
β-Hydroxydecanoyl Thioester Dehydratase, 260
γ-Aminobutyrate Aminotransferase, 262
Kinetics of Slow-Binding and Tight-Binding Inhibition, 268
Slow Binding, 268
Tight Binding, 269
Slow-Binding Inhibition, 270
Dihydrofolate Reductase, 271
Prostaglandin H Synthase, 274
Tight-Binding Inhibition, 280
HMG-CoA Reductase, 280
Alanine Racemase, 285
5-Enolpyruvoylshikimate-3-Phosphate Synthase, 289
Acetylcholinesterase, 291
Chapter 6. Acyl Group Transfer: Proteases and Esterases, 297
Chemistry of Acyl Transfer, 297
Serine Proteases, 300
Chymotrypsin, 301
Subtilisin, 311
Cysteine Proteases, 314
Papain, 315
Caspases, 317
Aspartic Proteases, 317
Molecular Properties, 318
Mechanism of Action, 320
Metalloproteases, 323
Carboxypeptidase A, 324
Thermolysin, 327
Esterases, 328
Structure and Function, 328
Phospholipase A2, 329
Contents xv
Chapter 7. Isomerization, 333
Aldose and Ketose Isomerases, 333
Chemistry, 333
Phosphoglucose Isomerase, 334
Triosephosphate Isomerase, 335
Xylose Isomerase, 341
Phosphomutases, 341
α-Phosphoglucomutase, 341
β-Phosphoglucomutase, 343
Phosphoglycerate Mutases, 343
Racemases and Epimerases, 346
Proline Racemase, 346
Glutamate Racemase, 350
Mandelate Racemase, 352
UDP-Galactose 4-Epimerase, 355
Ribulose-5-P 4-Epimerase, 360
UDP-N-Acetylglucosamine-2-Epimerase, 361
Chorismate Mutase, 364
∆5-3-Ketosteroid Isomerase, 366
Radical Isomerizations, 368
Glutamate Mutase, 369
Methylmalonyl CoA Mutase, 371
Lysine 2,3-Aminomutase, 376
Newer Isomerases, 379
UDP-Galactopyranose Mutase, 379
Pseudouridine Synthase, 379
Chapter 8. Decarboxylation and Carboxylation, 387
Chemistry of Decarboxylation and Carboxylation, 387
Decarboxylases, 388
Pyruvate Decarboxylase, 389
Amino Acid Decarboxylases, 394
Acetoacetate Decarboxylase, 403
Mevalonate Pyrophosphate Decarboxylase, 405
Radical-Based Decarboxylases, 407
Orotidine Monophosphate Decarboxylase, 414
Carboxylases, 418
Ribulose-1,5-Bisphosphate Carboxylase, 419
Phosphoenolpyruvate Carboxylase, 425
Vitamin K–Dependent Carboxylase, 426
Chapter 9. Addition and Elimination, 433
α,β-Elimination/Addition Reactions, 433
Cofactor-Independent α,β-Elimination/Addition Reactions, 434
Cofactor-Dependent α,β-Elimination/Addition Reactions, 440
xvi Contents
β,α-Elimination/Addition Reactions, 456
Methylidene Imidazolone–Dependent Elimination and Addition, 456
Carbonic Anhydrase, 462
Isomerization and Elimination, 465
Catalytic Process, 465
Coenzyme B12–Dependent Elimination, 466
Chapter 10. Phosphotransfer and Nucleotidyltransfer, 476
Chemistry of Phosphoryl Group Transfer, 476
Phosphomonoesters, 476
Phosphodiesters, 483
Phosphotriesters, 483
Five-Member Ring Phosphoesters, 484
Enzymatic Phosphoryl Group Transfer, 487
Single and Double Displacements, 487
Phosphotransferases, 489
Protein Phosphorylation: Protein Kinase A, 502
Phosphomonoesterases, 509
Enzymatic Nucleotidyl Group Transfer, 521
Nucleotidyltransferases, 521
Phosphodiesterases, 539
Chapter 11. ATP-Dependent Synthetases and Ligases, 547
Ligation and the Energy of ATP, 547
Activation by Phosphorylation, 548
Glutamine Synthetase, 548
Carbamoyl Phosphate Synthetase, 554
Activation by Adenylylation, 559
DNA Ligase, 559
Aminoacyl-tRNA Synthetases, 561
Ubiquitin, 566
Chapter 12. Glycosyl Group Transferases, 569
Chemical Mechanisms, 570
Chemistry of Glycoside Hydrolysis, 570
Enzymatic Glycosyl Transfer, 573
Glycosyltransferases, 575
Sucrose Phosphorylase , 575
Glycogen Phosphorylase, 577
Purine Nucleoside Phosphorylase, 584
Glycosidases, 587
Families and Structures, 587
Lysozyme, 589
T4 Lysozyme, 595
Contents xvii
Chapter 13. Nitrogen and Sulfur Transferases, 597
Nitrogen Transfer, 597
Aspartate Aminotransferase, 597
Tyrosine 2,3-Aminomutase, 602
Amidotransfer, 604
Glutamine:PRPP Amidotransferase, 607
Sulfur Transfer, 609
Biotin Synthase, 611
Lipoyl Synthase, 612
Chapter 14. Carbon-Carbon Condensation and Cleavage, 617
Chemistry, 617
Enolization of Acetyl CoA, 619
Acetyl CoA in Ester Condensations, 619
Citrate Synthase, 620
Thiolases, 627
Carbanionic Mechanisms, 630
Transaldolase, 631
Transketolase, 634
Serine Hydroxymethyltransferase, 639
Carbocationic Mechanisms, 645
Farnesyl Pyrophosphate Synthase, 645
Squalene Synthase, 648
Chapter 15. Alkyltransferases, 655
Chemistry of Alkylation, 655
Biological Alkylations, 655
Alkylation Mechanisms, 656
Enzymatic Alkylation, 657
Protein Farnesyltransferase, 657
Catechol O-Methyltransferase, 661
S-Adenosylmethionine Synthetase, 665
Methionine Synthases, 670
Chapter 16. Oxidoreductases, 679
Pyridine Nucleotide–Dependent Dehydrogenases, 680
Alcohol Dehydrogenase, 680
Lactate Dehydrogenase, 686
Short-Chain Alcohol Dehydrogenases, 687
Glyceraldehyde-3-P Dehydrogenase, 690
Glutamate Dehydrogenase, 693
Disulfide Oxidoreductases, 694
Dihydrolipoyl Dehydrogenase, 694
Ribonucleotide Reductases, 698
Classes of Ribonucleotide Reductases, 700
Structural Relationships of Ribonucleotide Reductases, 705
xviii Contents
Chapter 17. Oxidases and Oxygenases, 710
Oxidases, 710
D-Amino Acid Oxidase, 710
Monoamine Oxidases, 716
Isopenicillin-N Synthase, 718
Urate Oxidase, 721
Monooxygenases, 722
Lactate Monooxygenase, 722
Cytochrome P450 Monooxygenases, 722
Iron-Methane Monooxygenase, 727
α-Ketoglutarate–Dependent Oxygenases, 732
Dopamine β-Monooxygenase, 735
Copper-Methane Monooxygenase, 737
Nitric Oxide Synthase, 738
Dioxygenases, 741
Intradiol Dioxygenases, 741
Extradiol Dioxygenases, 744
Chapter 18. Complex Enzymes, 749
Multienzyme Complexes, 750
α-Ketoacid Dehydrogenase Complexes, 750
Pyruvate Dehydrogenase Complex, 750
Fatty Acid Synthesis, 757
Acetyl CoA Carboxylase, 757
Fatty Acid Synthases, 761
Modular Enzymes, 763
Polyketide Synthases, 763
Nonribosomal Polypeptide Synthetases, 767
Ribosomal Protein Synthesis, 768
RNA Polymerase, 768
The Ribosome, 770
Energy-Coupling Enzymes, 777
Nitrogenase, 777
Cytochrome c Oxidase, 782
ATP Synthase, 786
Myosin and Muscle Contraction, 792
Appendices, 803
Appendix A: Haldane Relationships for Some Kinetic Mechanisms, 803
Appendix B: Inhibition Patterns for Three-Substrate Kinetic
Mechanisms, 804
Appendix C: Equations for Number of Occupied Sites in the
Binding of a Ligand to a Multisite Macromolecule, 804
Appendix D: Derivation of Steady-State Kinetic Equations by the
King-Altman Method, 805
Index, 809
Enzymatic Reaction
Mechanisms
Other documents randomly have
different content
ruled Odin, and there too was a great place of sacrifice. Twelve
pontiffs (hofgodar) presided in the temples, who were at the same
time the judges of the law.”[50]
Defective as was the geographical knowledge of Snorro, he has,
no doubt, correctly assigned the cradle of this people, and of the
Vanir. They were neighbours; they were consequently often at war,
until the chiefs of both agreed, not only to be for ever amicable, and
to join in all future conquests, but in some degree to amalgamate by
a union of government. Hence the junction of the Vanir to the Aser,
and the contiguity of their respective regions in the Scandinavian
calendar. How Asgard and Vanaheim came to be placed in heaven,
as well as on earth, has puzzled many writers. They may be equally
puzzled, that the twelve drothmen, or pontiff-chiefs, should be
transfused into so many divinities; and the temple of the earthly
transferred to the celestial Asgard. There are two ways of solving
this problem. It is possible—it is even exceedingly probable—that the
Scythians, long prior to their migration from Asia, called their
country after the heavenly one which they expected to inhabit after
death. The government of the Aser was essentially theocratic, and
assimilated as much as possible to that which they believed to exist
above. Nor were they peculiar in this economy: Athens and greater
nations have done the same. The twelve great priests of Egypt were
named after the twelve gods who ruled the same number of celestial
signs. Such was the case in Assyria. In Persia, too, the number of
priests in the great temple corresponded with that of the
Amshaspands, or celestial genii, who governed the world as
vicegerents of Ormusd. Nothing, indeed, is more natural than the
position, that men devoted to the service of the gods would
endeavour to form their establishments after the model which the
gods themselves were believed to have adopted. “Thus, the Aser
were the gods of the new religion introduced by Odin, and at the
same time his temporal companions and followers,—the tribe of the
Ases, or Aso-Goths, from the river Tanais. Asgard, or Godheim, is
their celestial abode, from which they descended on earth
(Manheim) to mingle with the children of men; and is, at the same
time, the original seat of Odin and his people on the river Tanais.”[51]
This we consider the more natural solution of the problem in
question. It may, however, be, that the disciples of the original
pontiff began after his death to invest both him and his companions
with the ensigns of divinity, and assimilated them, both in number
and in attributes, with the ancient divinities of Scythia; making,
however, some change. In either case there must have been a
change. We have before expressed our opinion that Thor was not a
Scythian god: he, therefore, (and the same may be said of one or
two others,) must have been subsequently admitted into the divine
college, when the union for which we have contended took place
between the native and foreign religion; or rather, when the foreign
was engrafted on the native system. That system, we repeat, was, in
our opinion, the basis of the one contained in the Eddas; and much
more than the basis.
The union which we have endeavoured to establish, will account
for the elaborate, however heterogeneous, system of the Eddas.
That system was, assuredly, not the work of one people, or, we may
add, of one age. It was derived from people widely different in
character, habits, opinions, and manners; and it was probably the
work of centuries. The successors of the twelve original pontiffs
effected, there is reason to think, much more than they did, or than
their predecessors had done. The elements were, indeed, strewed in
Norway; but they could scarcely be fashioned into a whole; still less
could they have assumed that stately form which they exhibited in
the age of Sæmund and Snorro. They consisted of detached
portions, composed at different periods, and probably not connected
—not fashioned into a whole—until many centuries after Odin’s
death. Nay, there is some reason for concluding, that the two
Icelanders we have just mentioned were the first collectors of these
scattered fragments, no less than of the comments on each by the
recent Scalds of their own country, and the more ancient Scalds of
Norway. Of the same opinion the reader will probably be, before he
closes the present volume.
Having now given a general view both of the Scandinavian
universe and of its inhabitants, and shown the probable relation
between its gods and its mortals, we proceed, in the following
section, to examine these gods more in detail, and, where
practicable, to explain their respective attributes by the physical
phenomena on which they were so frequently based.
SECTION II.
CHIEF MYTHOLOGICAL PERSONAGES OF SCANDINAVIA.
ODIN, THOR, AND LOKE.—THEIR CHARACTERS PHYSICALLY INTERPRETED.—
THEIR WIVES AND OFFSPRING.—THE THREE DEMON CHILDREN OF LOKE.—
INFLUENCE OF THIS DEITY OVER THE FATE OF THE UNIVERSE.—HE IS
PRESENT IN EVERY GREAT MYTHOS.—RAPE OF IDUNA.—THOR’S VISITS TO
JOTUNHEIM.—THOR AND THE GIANT HYMIR.—THOR AND THE GIANT THRYM.
—NIVOD, FREYR, FREYA.—EXPEDITION OF SKIRNIR-ÆGIR AND RAN.—OTHER
DEITIES.—BALDER.—PUNISHMENT OF LOKE.—RAGNAROK.—RECOGNITION OF
A GREAT FIRST CAUSE BY THE PAGAN SCANDINAVIANS.
ODIN, THOR, LOKE.
The first two gods we place together, as well for the purpose of
comparison as that of contrast; the last, because his agency is
necessary to explain the other two.
According to the Eddas, Odin had several wives; the first was Frea,
or Frigga, by whom he had five sons, Thor, Balder, Braga, Hermod,
and Tyr: the second was Skada, by whom he had Semming; the
third was Grydur, by whom he had Vidar; the fourth was Rinda, by
whom he had Balder, or Bo.[52]
In Valhalla he has a table separate from the Einheriar, but he lives
only on wine; and the meats set before him he distributes to two
wolves which stand by his side. These are Geri the devouring, and
Freki the fierce.
He learns all that passes on earth, without the trouble of
ascending Lidskialf[53], by means of two ravens, which leave Asgard
at daybreak, and at dinner time return, to perch on his shoulders,
and whisper into his ear all that they have seen. These ravens are—
Observation and Memory; both presents from the enchantress
Hulda. Hence he is called the raven god.
These mythi are for the most part sufficiently obvious. Frigga is a
personification of the earth; while Odin himself, in his character of
chief god, may represent heaven. Heaven and earth give origin to—
thunder (Thor), the summer-sun (Balder), the swift messenger
(Hermod), the hospitable board (Braga), and the undaunted
defender of nature (Tyr). Skada, the daughter of the giant Thiasse,
and a nymph of the mountains, is a personification of the spring
winds; but we cannot see the import of the mythos—if, indeed, there
were any intended—in regard to Semming. Why Vidar should be the
offspring of Grydur is equally dark; but there is propriety enough in
making the frost (Rinda) the mother of barrenness. The two wolves
at Odin’s side denote his ferocity as the god of battles; and the two
ravens, memory and observation, explain his knowledge of the past
and present. To that of the future this god had no pretensions; this
was reserved to the Norny Skulda, and to a few of the Valas, or
prophetesses.
Immediately dependent on Odin—the ministers of his will as the
god of war—were the three Valkyrs, or choosers of the slain. They
also administer to the slain at his banquet.
There are many legends respecting Odin, who often visited
mankind. We select one because it illustrates the observations we
have made in regard to the rivalry of him and Thor. Sterkodder, the
celebrated champion[54], when a child, was taken captive. He fell to
the lot of one named Granè, or Whiskers, who was named Horsehair
Whiskers, and who brought him up as a foster-son. This was no
other than Odin in disguise, whose attachment to one destined to
become so unrivalled in arms may be easily conceived. One night
the destiny of the young man, unknown to himself, was to be shown
him. Horsehair Whiskers, of whose quality he was ignorant,
embarked with him in a small boat, and they proceeded to an island,
landed, and by midnight reached an open plain in the centre of a
forest. There he saw a large assemblage; and within the ring formed
by the assemblage were twelve seats, for so many judges. Eleven
were full, but one was empty, and Horsehair Whiskers immediately
seated himself in it. From the instantaneous salute of Odin by the
judges, and the mention of his name, the chieftain perceived that he
was in the awful presence of that deity, and of the other gods. Odin
said that the judges should now decree Sterkodder his destiny. Thor
then spoke and said, “Alfhild, the mother of Sterkodder’s father,
chose for her son’s-father (husband) a very wise Jötunn (giant) in
preference to Asathor; wherefore I appoint to Sterkodder that he
shall have neither son nor daughter, and thus be the last of his race.”
Odin. “I grant him to live three men’s age.”
Thor. “He shall do a vile act in each of them.”
Odin. “I give him that he shall own the best weapons and
harness.”
Thor. “And I appoint him that he shall own neither land nor sand.”
Odin. “I give him that he shall be rich in money.”
Thor. “I lay on him that he shall never seem to have enough.”
Odin. “I give him victory and martial skill in every fight.”
Thor. “I lay on him that in every fight he shall lose a limb.”
Odin. “I give him the poet’s faculty, so that he shall produce
poems with as much ease as unmeasured language.”
Thor. “He shall never be able to remember the verses he makes.”
Odin. “I grant him that he shall be favoured by those of greatest
rank and name.”
Thor. “He shall be hated by all others.”
Then the judges ratified to Sterkodder all that had been said, and
the council broke up.
The Saga from which this incident has been derived was written
by a Norwegian, who certainly held Thor to be the equal, if not the
superior, of Odin. It is not unlike the magian scene at the creation of
the world, when to every good thing decreed by Ormusd, an evil one
was joined by Ahriman. That Odin and Thor were rival deities, and
that they gave rise to hostile sects, is evident. And there is another
point from which this hostility may be viewed. The warriors who
went to Valhalla were all of noble birth; they were jarls or herser,
were rich and powerful. But what became of meaner freemen and
thralls (serfs) who fell in battle? They went to Thrudheim to the
palace of Thor, Bilskirner[55], which that the owner might not be
outdone, had the same number of gates as the palace of Odin, viz.
five hundred and forty. Does not this prove that Thor was the native,
Odin the foreign, god?—that the former belonged to the vanquished,
the latter to the victorious people? The very name of Thor shows
that he was a Celtic divinity. He is the Taranis of Lucan, the Toron of
the Scottish Highlands, and the Tiermes of the Lapps.[56]
The visit of Odin to the giant Vafthrudnir, and his contest with
him[57], may also serve to illustrate his boasted knowledge, as well as
power. Frigga, his wife, was alarmed when she first heard of her
husband’s intention to visit “that learned giant.” He conquers,
indeed, in the strife, but not through any superiority of knowledge: it
is rather by an unworthy artifice.
Frigga, the wife of Odin, was a distinguished personage in the
northern Olympus. She is the queen and mother of the gods. Her
palace, called Fensale, was magnificent; and it was a sort of
drawing-room for all the goddesses. Her prescience was great; she
could foresee the future, and she was invoked by women in
childbed.
According to the vulgar genealogy,—that which the Odinists, in
opposition to the Thorists, were anxious to establish,—Thor was the
eldest son of Odin and Frigga. Even in Sweden he was, after Odin,
the first in rank among the gods. We may even doubt whether by
one sect of the Odinists he was not esteemed the first; for his image
at Upsal, where he is represented seated on a throne, with the
attributes of divine majesty about him,—while Odin, the war god, is
standing at his right hand with a drawn sword, and Frigga, the
goddess of production, on his left, with the fruits of nature in her
hands,—clearly establishes his predominancy. His strength was
unrivalled; and his structure so large, that no horse could carry him:
he always travelled in a chariot drawn by two he-goats. He had three
treasures, all unrivalled, all made by the Dwarfs. Of these the most
famous was his hammer, called Miölner (the miller, the bruiser),
which, when thrown by his powerful hand, was irresistible; yet,
however far it was thrown, it always returned to him. Formidable as
it was, it was so small that he could put it in his pocket. No hands
but his could touch it; nor even he without his wonderful steel
gloves, the second of his treasures. The third was a belt,—
Melgingandur, which doubled his strength whenever he girded it on.
Above all the gods, he was the enemy of the Rimthurser, or Frost
Giants, against whom, with his dreaded weapon, he waged
unceasing war. The very glare of his eyes was tremendous: it was
lightning; and lightning was emitted by his chariot wheels as he
rolled along. Every day did he make the circuit of Asgard, to drive
away the giants.
Of this mythos an interpretation is scarcely necessary. Miölner is
his thunderbolt. His antipathy to the giants—the powers alike of
darkness and of cold, and his daily circuit round Asgard, sufficiently
explain themselves. His gloves and belt were an embellishment,
which have no necessary connection with his nature. The latter is to
be found in many oriental fictions, (the Arabian Nights, for instance,)
and in many also current throughout Europe. His wife, Sif, is another
illustration of the mythos. She is held to be a personification of the
summer earth, and is represented in the act of distributing fruits and
flowers. She, like her husband, was peculiarly worshipped in Norway.
By a former husband she had a son—Uller, the god of hunters,
whose residence was Ydale, or the Dewy Valley.[58] The most
wonderful of her peculiarities was her hair, which was unrivalled for
its beauty, and to which we have before adverted.
The fact, that Sif was worshipped in Norway alone, of all the
Scandinavian regions, is another argument in favour of her
husband’s supreme worship, long before the arrival of the Aser. A
still stronger one is to be found in the fact, that Thrudheim, or
Thrudvang, was the name of a district in that kingdom, no less than
of a palace in heaven: and the strongest of all is, the peculiar
affection with which he was regarded by the Norwegians, who held
him to be their native, their tutelary god. He seems to have had
some attributes of the Roman thunderer: the same day (Thursday),
and the same planet (Jupiter), were sacred to him.
The giants of whom Thor was thus the natural, the everlasting
enemy, were, as we have frequently observed, the offspring of
Bergelmer, the old man of the mountains, and of his wife, who
escaped the destruction of their race by the blood of Ymer, only
because they chanced to be at sea, fishing, when the giant was
slain. Repairing to the dark lower region which lies within the polar
seas, they soon peopled it. Darkness, indeed, was the element of
these beings: no sun enlightened or cheered them. When they
visited earth, it was during the night, for then their power was the
greatest. In magic they surpassed all other beings: they possessed
many secrets, relating to the origin and nature of things, unknown
to the wisest of the gods. With them the three Nornies, or destinies,
—with them Vala herself, the great prophetess of heaven, was
educated. They regarded the Aser with dislike,—as usurpers of a
world which rightly belonged to them; and towards the sons of
Askur, the creation of the gods, they bore equal dislike. This feeling,
indeed, did not prevent the Aser from occasionally intermarrying
with them; but the marriages were never well assorted. The king of
this vast gloomy region was Ugarthiloc, or, more correctly,
Utgardelok, viz. the Loke of Utgard, the monarch of the outer world.
The notion entertained of this personage, and of the whole race, by
the Danes, we have shown on a former occasion.[59] Wild as the
legends there related may seem, they have their meaning. The
reader will not fail to observe, that these original inhabitants of the
earth—this people destroyed by the Aser, and exiled into the dreary
wastes of the North, were the original Finnish, or rather Celtic race,
whom the Goths expelled. The mythology of that race was full of
giants; the Druids boasted of an acquaintance with nature denied to
the rest of mankind; and the boast was probably a just one. The
testimony borne by Cæsar to the extensive character of their
knowledge, will abundantly illustrate this part of the historical
question. Again, the Celts pretended to mystical science: in proof of
it, look to Cæsar, to the traditions rife wherever the Celts have been
located, and, above all, to the fragments of the ancient Welsh bards
preserved in the Archæologia of the principality. The Eddas are filled
with Celtic mythological allusions. For example, Celtic were the
dwarfs or fairies of the benevolent class; while the malignant ones,
who were a kind of evil genii, came with the Aser from a seat where
the two principles of good and evil were a dominant article of the
popular creed.
A personage no less important than Odin or Thor in the
Scandinavian mythology, is Loke, or, as he is sometimes called,
Luptur. He was important, not from his power, or his wisdom, or his
dignity, but from his cunning, his treachery, his ill-nature, and the
influence which he exercised alike over gods and men. He was the
son of the giant Farbautè, by the enchantress Laufeya. Though of
giant race, he obtained admission among the gods: indeed, as his
manners were exceedingly pleasant, his mirth constant, and his wit
unbounded, whenever they were not mixed with spite, he could not
fail to be acceptable to so vulgar a race as the Aser. But when, as
indeed was often the case, there was malice in his jokes, his
laughter made the hearer shudder. Why the gods should tolerate
him, is not very clear; but destiny was probably the reason which a
devout Odinist would have assigned for it,—a very convenient reason
in most systems of mythology. His birth might be traced to the origin
of time; for, in some way or other, he was concerned with Odin in
the work of creation, though the connection is very obscurely hinted
at. He was a relation, we are told, of the Utgard Loke, or Ugarthiloc,
the monarch of the frosty giants. These two personages were no
doubt originally the same; but as the Celts and Aser had different
notions of the same being, it was found necessary to introduce the
two into the united creed. In virtue of his connection with them,
Loke often visited the giants, by whom he was as little trusted as by
the Aser. But he was sometimes useful to both; and, from the malice
of his nature, no less than from his dislike to the gods, whom he at
once feared and hated, he was frequently the ally of the giants in
their efforts to recover their lost dominion, and to destroy the
usurpers. If he thus brought the latter into danger, he alone could
extricate them from it. In perfect accordance with the Eddas, he is
thus described by Ohlenschlager:—
Amongst bright Asgard’s lords
Is one, As-Luptur hight.
Like honey are his words;
His heart is filled with spite:
His form is passing fair,
And winning is his mien;
But still his guileful leer
Shows all is false within.
Though oft his traitorous wiles
The Aser’s wrath provoke,
His smooth tongue still beguiles,
And stops the impending stroke.
Oft cited to appear
He cowers the Ash before.[60]
At Odin’s table near
His place to Asa Thor.[61]
He was, indeed, as a god, the familiar companion of Thor; who,
however, had no great wish for his society. Like most of the gods, he
was married. His wife, Signe, was an amiable being, who loved him
in spite of his depravity. By her he had two sons, Nari and Vali,
whose fate will be mentioned in the proper place. But he had other
and more mischievous offspring by the giantess Augerbode,—Fenris
the wolf, Jormungandur the great serpent, and Hela the queen of
death. This alleged affinity will confirm the observation, that there
was originally but one Loke, the lord of Utgard, and consequently
the everlasting foe of the gods. How the Asgard Loke should become
so wicked as to produce such offspring, might surprise us, if we
were not assured that he was not so originally, and that he became
so by eating the half-roasted heart of an enchantress.
These three children of Loke were reared in Utgard by the mother.
The fatal influence which they were to exercise over the universe,
was not concealed from Vala, the mysterious prophetess of heaven,
or from Skulda, the Norny of the future. The gods being warned,
sent to secure them. Jormungandur, one of the most dreaded, was
seized, and by Odin cast into the great sea that separates the
human from the giant world. There so large did it become, that it
surrounded the whole earth,—being condemned to hold its tail in its
mouth, and thus to form a circle. There he lies, waiting for the time
when destiny will unloose him—the Ragnarok, or the twilight of the
gods; when he will assist in the destruction of the visible universe.
Hela, the next mythologic offspring, is hideous to behold,—her
body being half livid, half of natural colour. By Odin, or rather by
destiny, of which he was merely the instrument, she was placed in
the upper confines of Nifleheim,—in the region which, from her, is
called Hell (Helheim). She was invested with dominion over six, or
perhaps seven, of the nine worlds, (as we have before observed,
there is some doubt whether Muspelheim be eternal,)—over men,
and dwarfs, and giants, and gods. All who die a natural death
proceed to her “drear abode:” hence her title, queen of the dead.
“Hela’s hall,” says the prose Edda, “is affliction; her table is famine;
her knife is hunger; her threshold, a drawbridge; her bed, lingering
sickness; her tent, cursing.” She too, like Odin, had nornies, whose
province it was to summon mortals to her vast domain. But these
were much inferior in loveliness and dignity to the celestial nornies.
They appeared to the fated victim by night only. Hela herself was
sometimes believed thus to appear. She had a dark red cock, to
signify, by its crowing, the approach of fate; and a spectre horse, to
carry the doomed to her gloomy abode.
The third demon offspring of Loke, the wolf Fenris, is no less
wonderful than his brother and sister. The one had been surprised
and, thrown into the sea; the other had been partly persuaded to
submit, through the high dignity offered to her; but Fenris, who was
more powerful, was also more troublesome. He was taken, indeed,
and bound; but he snapped his fetters, strong as they were, as if
they had been nothing. A massive chain was now made, and he was
bid to try its strength: it snapped as if it had been dried clay.
Another was made double the strength of the preceding,—the
strongest that the gods could make; but with a very slight effort it
too gave way. What was to be done with this formidable criminal,—
one destined, if oracles were true, to endanger the world? The gods
had no fetter in which to bind him; the giants, who were skilful,
could not be expected to join in any design against one of their own
body,—one, too, that was naturally hostile to the Aser. In this, as in
many other dilemmas, recourse was had to the Dwarfs in the bowels
of the earth. At the instance of Skirnir, the messenger of Freyr, they
constructed a chain called Gleipner; which, though so slender as to
resemble a silken thread, was nevertheless not to be broken by
gods, or giants, or dwarfs. The Edda acquaints us with the materials
of which it was constructed. These were six, all curious enough to
deserve mentioning:—the sound made by the feet of a cat; the
beard of a woman; the roots of huge rocks; the fibres of trees; the
breath of fishes; the spittle of birds. But how bind by it the
formidable monster? Deceit must be used. Repairing with him to a
solitary island, the gods desired him to try his strength on this, as he
had done on preceding things. “Little honour,” replied the cunning
demon, “can result from breaking a silken thread; but probably it
may be enchanted!” and he refused to try it. He was next taunted
and jeered; and in vexation he at length consented to be bound; but
then, to be assured that the gods were honest in their proffer, he
insisted that some one of them should put a hand in his mouth.
They were in utter dismay; but the undaunted Tyr[62], the northern
Mars, the defender of the gods, at length resolved to sacrifice a
member for the preservation of the universe. He therefore placed his
hand in the open jaw, and the wolf allowed himself to be fettered.
The chain was cunningly fastened round his body, passed through a
rent rock, carried downwards to the centre of the earth, and there
made fast. Fenris now tried as before; but so far from escaping,
every effort that he made only entangled him the more, and rivetted
his bonds the more firmly. He therefore desisted; but in his anger he
bit off the hand of Tyr. From that moment the god has been only
left-handed; but as he uses that hand with much effect, he is still to
be dreaded. He alone had courage to take food to an animal, the
roaring of which was felt by all nature, until the gods thrust a sword
into his jaws, and thus gagged him. There he lies until Ragnarok,
when, like Midgard’s serpent, he will break loose.
There is no personage in the whole system of a more mythic
character than Loke. He was evidently the personification of the
active evil principle. His name signifies flame; and he is a
representative of the demon of fire—the destructive, in opposition to
the alimentary, aerial fire, of which Balder may be considered the
symbol. At this day the devil is called Loke by the Norwegians. Still
there is frequently some obscurity in the mythi respecting him, and it
is occasioned by his being so often confounded with the demon king
of Utgard. Though they were originally one, the Edda has made him
into two, in conformity no doubt with the genius of two distinct
systems of mythology. The mysterious allusion to the assistance
which he afforded Odin in the work of creation, is one great proof of
his identification with the powers of evil: his relationship with the
giants, on both sides, sufficiently accounts for his hostility towards
the gods, with whom he associated that he might find an
opportunity of triumphing over them. He is styled a coward, because
his deeds will not bear the light—the inventor of deceit, of lies, of
every thing base. The first of his offspring, the great serpent, is
evidently a relic of the Celtic creed. The Britons acknowledged its
existence; and there are two bold promontories on the coast to
which they have given the name of the Worm’s Head.[63] Of the wolf
Fenris the character is more obscure, though no less confirmatory of
the mythos. It is doubtless a symbol of destruction. In several
countries of the East, it is believed that a wolf will finally destroy, if
not the world, the sun and moon. Thus, in the Budhist system, a
wolf, Rakoo, is always on the watch to swallow both luminaries. This
mythos, we suspect, with a living writer[64], has given rise to the
superstition so common in the middle ages,—that of men-wolves;
viz., the power possessed by some men of assuming the form of that
animal. Hela, or death, the offspring of sin, or Satan (Loke), needs
no explanation. We may, however, observe, that there is some
plausibility in the arguments of Magnussen, when he attempts to
show that Helheim is more ancient than Valhalla; that it is the place
of punishment acknowledged by the original inhabitants, while the
warrior’s heaven was introduced by the Gothic conquerors.
The mythological fables in which Loke so prominently appears, will
illustrate his character better, and certainly more agreeably, than any
formal description. In most of them he was associated with Thor;
but we select one in which Odin and Hoenir were concerned with
him. Hoenir, we must observe, is but another name for Vile, the
brother of Odin, who assisted in the work of creation.[65]
RAPE OF IDUNA.
The three Aser one day left Asgard to see other worlds, especially
Utgard. Travelling over dreary wastes, they reached a mountainous
region, more hungry than they had for some time been. Entering a
valley, they found a herd of cattle, and killed one of the animals for
supper. Loke, who was to be the cook, made a fire, and proceeded
to his task, while the two nobler gods walked about. But
notwithstanding the great heat of the fire, the ox would not roast. A
voice, from the tree above him, told him that he would have no
supper unless he promised to let the speaker join. He looked up, and
seeing an eagle only, gave his consent. The bird now descended to
the fire, and seized both shoulders, which he, considering as
somewhat too large a share, would not permit. Taking a large billet
of wood, he struck the unreasonable animal; when the eagle
instantly flew upwards, one end of the billet adhering to its beak.
But alas! the other end was no less tenacious of Loke’s hand; and
away he was dragged over mountain, wood, and stream, his arm
ready to fall from his body, and his feet sorely wounded by being
trailed over the sharp rocks and bushes. He lustily called for help to
Odin and Thor. “Cry away!” replied the eagle, who was no other than
the giant Thiasse in that shape[66]; “but never shalt thou be released
from this situation, unless thou promise by oath to bring Iduna and
her apples from Asgard to me!”
Iduna was the wife of Braga, the god of eloquence, and daughter
to the dwarf Ivalldr, one of the most scientific of his race. She was a
goddess, and the wife of a god: for both honours she was no doubt
indebted to the wonderful apples of which she was the guardian,
and which had been given her by her kindred. They had this virtue,
that when the gods felt the approach of age, they had only to eat of
these apples to be restored to all the bloom of youth. The giants,
like the gods, were subject to decay; and, like the gods, they wished
for the means of immortality,—to escape the dark empire of Hela.
As Loke was no friend to the Aser, he swore to comply with the
giant’s demand, within a given time. He was therefore released, and
enabled to return with the two gods to Asgard. When the
covenanted time arrived, he told Iduna, that in a neighbouring wood
he had discovered some apples, much finer, and much more
valuable, than any she possessed. Her curiosity being raised, she
took some of her own apples with her, to compare with the others,
and was accompanied by Loke to the wood; but scarcely had they
passed the boundary of Asgard, than Thiasse arriving in the eagle’s
shape, bore her away to the dark mountains of Utgard.
Great was the consternation of the Aser at the disappearance of
Iduna and her apples. The effect was soon visible: they became
weaker, less supple, decrepit, and wrinkled. Though the season was
spring, the flowers withered, and the leaves became sear as at the
close of autumn. A council of the gods was convoked to learn how
and whither Iduna had disappeared. No one could give them any
other information than this,—that she had been last seen with Loke
departing from Asgard. Loke was examined; and when he showed a
disposition to evade the questions that were asked, Thor seized him,
and threw him into the air so high that his heels struck the moon,
and then descended to the sea. All this was nothing in comparison
with what he would suffer if he did not restore the goddess. He
readily promised to do so, if Freya would lend him her disguise, that
of a hawk. Being furnished with it, he flew in that disguise to Utgard,
and reached the abode of Thiasse just as that giant had left it to row
for a short time on the neighbouring sea. Changing Iduna into a
swallow, he returned with her in his claws towards Asgard. When
Thiasse returned, and learned the departure of the goddess, he
resumed his eagle’s dress, and rapidly followed in the direction
which the hawk had taken. He obtained sight of the fugitives just as
they approached Asgard; and he would certainly have overtaken
them but for a stratagem of the Aser, who were anxiously watching
the pursuit. Forming a vast pile of faggots under the walls of the city,
they set fire to it; and the flames ascended so high as to burn the
eagle’s wings. Thiasse fell to the ground, and was immediately
despatched by Thor.
This is one of the most interesting fables of the prose Edda. It has
doubtless a meaning, though we are by no means sure that
Magnussen has discovered the right one. According to him, Iduna is
the spring, which may be called the renewer of nature’s youth.
Spring is always accompanied by joy and harmony,—by the song of
birds, by the cheerful hum of men, by the gambols of animals, by
the sportive winds: hence it is personified in Iduna: she is the wife
of Braga, the god of poetry, of music, of song, of harmony. Thiasse,
the giant, is the winter: Iduna flies from him in the shape of the
swallow, which is everywhere the bird of spring. The destruction of
the giant by the flames, denotes the season of winter killed by the
heat of the spring.—That this explanation of the mythos is ingenious,
as well as plausible, cannot be denied; but we are not quite satisfied
with it. Though a meaning is involved in these fables, we doubt
whether all the incidents are thus designed. Many were invented
through the love of invention, or rather to please the multitude; and
by such inventors physical principles would not always be observed.
For this obvious reason, much caution is requisite in interpretations
which have not positive authority for their base.
The next mythos in which Loke is exhibited, is in connection with
Thor.
THOR’S VISITS TO UTGARD.
Geyruth, also called Geirrod, was one of the Aser’s most
formidable enemies. In the former volume we have given, from Saxo
Grammaticus, a description of his empire[67],—a description rivalling
in power of invention any to be found in Homer. To it we refer the
reader, before he proceeds any farther with this narrative, as nothing
can be more curious than to compare the account which Saxo
derived from tradition (no Edda had then been compiled), and, what
is more, from Danish tradition, with that given in the sacred books of
the Scandinavians from Norwegian sources.
Thor’s first journey was preceded by that of Loke. Loke, with all
his cunning, was frequently in trouble;—and how could the devil be
otherwise? Assuming a hawk’s disguise, (the hawk in more countries
than the North was the symbol of that personage,) he entered the
dominions of Geyruth, was caught, and, when he refused to answer
the questions that were put to him, was shut up in a chest during
three months. His revenge then gave way, and he confessed who he
was. The giant then released him, on his promise to bring Thor to
Utgard, without belt or hammer. The object of the giant’s policy may
be easily guessed. Thor, the defender of Asgard, the everlasting
enemy of the giants, would be reduced to the same level with
themselves when deprived of those wonderful treasures. Loke had
no difficulty in prevailing on the stout-hearted god to visit the
dominions of the giant king. On the way to that region, within the
boundaries of Utgard, was a magic forest, of which the trees were
all iron. It was inhabited by certain enchantresses, who were the
mothers of male and female sorcerers, who could at any time
assume the wolf’s shape. These enchantresses were cruel: they
often raised storms, and enticed travellers into their power from the
mere love of destruction. Thor met one of these witches, who
cautioned him against the arts of Geyruth, and presented him with a
pair of iron gloves, a girdle, and a staff.
On reaching the river Vimur, the longest one in the world, he
observed Gialp, one of the giant’s daughters, standing astride the
whole river,—one foot on each bank; and making the water rise in a
fearful manner. He threw a rush at her, and forced her to retire.
Wading across, he proceeded to Geyruth’s palace, which he entered,
and a separate lodging was provided for him. In one corner of the
cavern was a stool, on which he sat down; but scarcely was he
seated, when the stool began to rise from the ground. With the staff
which he held, he struck the roof of the cave, and immediately heard
a loud scream beneath him. On looking, he discovered, with broken
backs, three daughters of his host, who had placed themselves on
the roof with the design of crushing him to death. Geyruth himself
did not escape more easily. Inviting Thor to drink with him, the two
sat down in another part of the palace, one on each side of a large
fire. Having sat for some time, the giant seized a red-hot iron wedge
that was glowing in the fire, and threw it with all his might at the
god. The latter caught it with his gloved hand, and returned it with
such force, that though Geyruth had run behind a pillar, it went
through both pillar, himself, and the walls of the palace. Still it
remained in his breast; and in that position, attended by his three
maimed daughters, he has remained ever since.[68]
In the second journey, which is much more imaginative, Thor was
accompanied by Loke. The temple of Upsal had been visited by
Utgardelok (the demon king of Utgard)[69], who had not only
extinguished the sacred fire, but made a ruin of the edifice. Now
Upsal was the palace which, above all others in Midgard, Odin loved.
In it Thor and Frigga too were worshipped with great pomp; and the
priestesses of the latter were of royal blood,—the daughters of
kings. Great was the wrath of the three deities. Thor, in particular,
was observed to knit his brows, and to clench his fist at table: but he
spoke not a word; for he was revolving the means of vengeance.
Formidable as he knew the demon king to be in natural, and still
more in supernatural power,—in a science unknown to the gods,—he
resolved to invade his dominions. Having emptied a full horn
presented to him by one of the Valkyrs, he called for his car, for his
goats, and for Loke, as the companion of his journey. Having
harnessed the animals, nailed on their golden shoes, wound the
reins round his waist, he entered with Loke; and grasping Miölner in
his right hand, proceeded at a rapid pace down the bridge Bifrost.
Adown the pointed way
As drove the impetuous god,
The red flames, lambent, play
Along the wheel-tracks broad.
Heimdal his horn blew loud,[70]
The god with sleepless eye;
Seven maids submissive bowed[71]
As the gold car flew by.
On earth some meteor dire
Men thought then to behold;
The heavens were fraught with fire;
In peals the thunder rolled.[72]
Reaching a cottage towards nightfall, they asked for hospitality,
which was readily granted. Humble was the cot; and it contained
little for gods to banquet on,—nothing but simple vegetables. But
Thor was not anxious on this account. With his hammer he slew his
two goats, which were skinned and roasted with considerable
despatch. Ample was the entertainment; not only was the flesh
delicious, but the place teemed with excellent mead; and some idea
may be formed of a divine appetite, when we add, that the two
goats were entirely devoured,—all but the bones, which Thor desired
should be carefully thrown back into the skins that were stretched
before the hearth. But Thialf, a son of the rustic host, and a mere
stripling, broke a thigh-bone of one goat for the sake of the marrow.
The next morning, before daybreak, Thor arose, and swung his
hammer over the two skins, when suddenly the two goats rose up as
if nothing had happened. But one of them limped; and dreadful was
the countenance of the god. Supplication, however, disarmed him;
he took the youth and a sister into his service; and leaving the car
and the goats at the peasant’s cottage, all four proceeded on foot.
The boy, who immediately won the favour of his master, carried a
wallet; and the maiden, quite a beauty, tripped lightly along. Thor
marched pensively; his hammer flung over his shoulder; his dark
locks escaping from his silvery casque. They reached the sea, which
was agitated by a dreadful tempest; and Loke began to be afraid;
but he was compelled to follow the god, who rushed into the water,
like some thundering rock. The mortals too followed: but the storm
continued to rage; and they required all the help of the leader to
reach the other side. A trackless desert was next to be traversed;
and on they went, in darkness, except that the moon now and then
gleamed,—weary, hungry, wet, and faint. Other trials were to be
encountered,—the storm, the lightning, the slippery ice, the deep
mud; the roaring wind, which the demon king excited by his magic
power. Thor, who had to support the maiden Roska, lost his temper;
and he vowed revenge on Utgardelok when he should meet him.
What seemed to be a hut, in the midst of the pitiless waste,
presented itself; and three of them entered it, Thor himself
remaining at the entrance with his mallet in his hand, to protect
them while they slept. Vast, and of a strange form, was the only
apartment which the hut contained; but in a storm any port is
welcome. Towards morning, while Thor glanced in great anger over
the waste, he heard a strange noise, and felt a strange motion.
Rising, he beheld, by the faint glimmer of the moon, a vast giant—so
vast as to cover several acres—asleep and snoring. Grasping his
mallet, he was preparing to punish the intruder, when up started the
giant. “Who are you?” demanded Thor. “Skrymner, the servant of
king Utgardelok, just come from Jotunheim.” He addressed Thor by
name, of whose feats he had heard much; but common report, he
thought, had been too favourable; for after all, even he, who was of
little esteem compared with his fellows, could put this hero of the
gods in the palm of his right hand. “I have lost my gauntlet!”
suddenly observed the giant, who groping for it, took it up. What
was it but the strange hut in which Loke and the two mortals had
passed the greater part of the night? All but Thor were dismayed at
this commencement of their acquaintance with the subjects of
Utgardelok; but Thor trusted in his hammer. “What brings you so far
to look at a desert?” was the natural question of Skrymner. Thor
replied, that he was determined to see, face to face, their boasted
monarch, whose magic and frozen mountains he only ridiculed. The
giant thought he might rue his boldness: however, if he was
determined to proceed, let him do so, and he (Skrymner) would be
his guide. When evening came, and the giant laid down to sleep
under a great tree, until supper was ready, there was more magic.
Neither Thialf, nor Loke, nor Thor himself, could open the wallet, or
cut the strings. In great wrath the god seized his hammer, and
struck at the forehead of the sleeping giant. “Has a leaf fallen on my
face?” asked the giant, rubbing his face, and wondering why they
had not gone to sleep. Towards midnight, the snoring of Skrymner
so enraged Thor, that he arose, and aimed a hard and more vigorous
blow at the monster: the hammer seemed to enter his very brain.
“Has an acorn fallen?” was the cool observation of the other, as he
rubbed his face. A third blow, which seemed to send the very handle
into the giant’s head, had no better effect; so that Thor now began
to have less confidence in the weapon which had hitherto terrified all
created things.
But we must not dwell on events which have been so frequently
described.[73] The adventures of the god and his companions at the
palace of Utgardelok were not such as to inspire him with more
confidence. Loke—fire itself, which consumes all things—was beat at
eating. Thialf—a mythologic personage too, though represented as a
peasant’s son, his name signifying thought—is exceeded in the race.
The mighty thunderer himself is vanquished in three successive
trials. Though he is the sun, the greatest drinker surely in all nature,
he cannot much lessen a large horn of liquor that is presented to
him: he cannot lift a huge tom-cat from the floor: he cannot, in
wrestling, throw a toothless old woman, who brings him on one
knee. In much shame, though in no consternation, the god returned
with his companions. On leaving the confines of the city, however, he
was made acquainted with the deceptions that had been practised
on him. The three blows which he had struck, were not at a head,
but at a rocky mountain; and deep were the dells which they had
made in it. The horn was the ocean; yet he had drunk so much of it
as to leave in many places land instead of water. The cat was
Midgard’s great serpent, which he had almost lifted from the sea.
The old woman was Hela, the goddess of death, who with all her
strength could only bring him on one knee. In great anger, he was
going to exact revenge for such tricks, when the spectre and the city
itself vanished like mist.
This mythos in a great degree explains itself. The contest between
Thor and Utgard’s monarch is evidently one between the summer
and the winter, between heat and cold, between light and darkness.
Many of the details, we believe, in opposition to Magnussen, who
sees in every thing a physical meaning, to have been created
without any other design than entertainment.
Hymis-quida, a song about Hymir, is from the elder Edda, and is of
great antiquity.
THOR AND THE GIANT HYMIR.
The sea-god Ægir gave a banquet to the gods; but he was little
prepared for such drinkers, and his mead fell short. Thor called for
more with some anger; and that anger was not diminished when he
found that no more was forthcoming. The excuse was, that Ægir had
not a cauldron large enough to brew sufficient mead at a time. Tyr,
who was present with the rest, and who, though a god, is
represented as the son of the giant Hymir, observed that his father
had one a mile deep, which might be obtained by stratagem. On this
business the two gods immediately departed in the chariot of Thor
towards Hymir’s abode, which lay on the confines of the eastern sea.
Here they found two ladies, the mother and wife of the giant; the
former a strange creature, with 900 heads; the latter, who was the
mother of Tyr, a fine woman, and kind as she was comely. She told
them both that she feared Hymir’s return; since he was subject to
dreadful passions; and she hid them behind some kettles. Towards
evening he returned, in no good humour. As he entered the house,
the icy mountains emitted a thundering noise. An old man he was to
view, and the hairs of his head, which resembled a forest, were
frozen. His wife, saluting him, told him that their son was arrived, in
company with the famous enemy of the giants and the friend of
men, Veor.[74] “Look,” she added, “where they sit, at the extremity of
the house, to avoid thy glance!” The giant looked; but they were
concealed by the nine kettles. At his glance, however, the tree or
beam from which they were suspended, burnt into two, and eight of
them burst. The two gods now advanced; and though Hymir was
compelled to exercise some degree of hospitality, he did so
unwillingly. Three oxen (one for each, we suppose, unless the lion’s
share was to be Hymir’s,) were ordered to be roasted. But Thor
showed that he had more than a giant’s appetite; for, to the surprise
of his host, he ate two of the animals himself. This made the latter
observe, that the next evening the two visitors must eat what they
could take in hunting or fishing for themselves. The next day,
therefore, Thor proposed to fish, if the giant would give him bait.
“Go amongst the cattle, and seize one,” was the reply. “I suspect,
however,” Hymir added, “that thou wilt not easily catch such bait.”
Without reply, Thor went into the wood, and seizing the horns of a
large black bull, pulled off its head, and returned to the giant, who
expressed some surprise at such a feat in one so little. They now
went out into the sea, and the giant hauled two whales. But nobler
was the prey of Thor: with the bull’s head he caught the great
serpent Jormungandur, the head of which he drew out of the water,
and which spewed venom upon him. The rocks trembled; the desert
places howled, and the ancient earth rolled itself closer. He then
struck the monster with his mallet, and it sank. Hymir rowed back,
sullen and silent; and the strength which had been exhibited in
bringing the two whales to his mountain home, gave him some
reason for thought. When returned, the two gods were desired to try
their strength in other things. A cup was put into the hands of Thor,
and he was defied to break it. In vain did he dash it against several
pillars in succession: he split them, but it remained unbroken. The
wife now whispered him to throw it against the giant’s head, which
was much harder than the rock. He did so; the head was uninjured;
but the cup was broken, and the owner lamented its loss. The next
trial of strength was to carry the great cauldron out of the house.
Tyr tried twice; but could not so much as move it. But Thor placed it
upon his head; and though the edges descended to his heels, he
walked away with it. He was now pursued by a great number of
giants whom he slew with his mallet.—From that time the sea-god
was able to treat the Aser men to their satisfaction.
Of this mythos the physical meaning is dark; and this darkness is
probably owing to the fondness with which the northern scalds
added extraneous circumstances for the sake of embellishment.
Nothing, indeed, is more hopeless than the attempt to restore these
ancient pieces to the original fragmentary state in which they were
left by the priests of Thor and of Odin. The scald has, by
embellishing, concealed the priest; the fabulist concealed the
philosophic theologian. All that we can safely assert is, that there is
here a physical contest between heat and cold, between evaporation
and congelation; that the sun (Thor), having drunk up all the
streams of the earth, now invades the dominions of frost and snow.
The bursting of the vessels under the glance of Hymir, is a notion
universally diffused in the northern latitudes. Thus, the two
magicians the suitors of Gunhilda, could destroy every thing by their
glance.[75] The meaning doubtless is, that excessive frost makes
every thing brittle, and may therefore be said to split every thing.
If the scalds took such liberties with the ancient or poetic Edda, as
often to bury the sun, they were more licentious still in regard to the
younger or prose Edda. This work was evidently compiled to explain
the former. With it a licence still more dangerous has been taken; so
that, in many instances, it bears little conformity with the preceding
work. We may add, that by modernising, paraphrasing, and
embellishing the prose Edda, Ohlenschlager has done no service to
the ancient mythology of his country: he cannot be followed by any
one that would form a correct notion of the subject. In the same
manner as the compilers of the second Edda deviated from the spirit
of the first, so has the celebrated Danish poet deviated from them.
[76]
For the sake of illustrating this divergence, let us advert for a
moment to the same adventure in the prose Edda and in the version
of Ohlenschlager: it will be found to have lost its mythical character
in proportion to the improvement of its fable.
When Thor reflected on the gross impositions which Utgardelok
had practised on him, he was apprehensive, and not without reason,
that gods and men would take him for a fool. To vindicate his merits,
he ventured again to visit Utgard, and without Loke, whose honour
he justly suspected. This time he would, like them, change his form,
and he obtained from Odin, in the shape of ointment, power that
would enable him to do so. Leaving behind his car and his goats—
O’er Dovre’s ridge[77] he strode,
For cliff nor torrent slack’d;
The tall pines, where he trode,
Like field of stubble crack’d.
Sneehattan’s peak of snow,
And Jotunfieldt he past,
Then sought the plains below,
And the sea reach’d at last;
He mark’d in curling wreath
The dull wave roll away,
And saw where, far beneath,
The serpent, brooding, lay.
His heart with hope beat high,
His voice shook as he spake,
Turning to Heaven his eye,
“No more, accursed snake,”
Quoth he: “in giant bend
Earth prison’d shalt thou keep,
Nor struggling sea-man send
To fell Ran’s cavern deep.”
“But being now resolved to proceed with caution, he began by
changing his form. Throwing his ponderous helmet on the ground, it
became a rock covered with pines.
Welcome to our website – the ideal destination for book lovers and
knowledge seekers. With a mission to inspire endlessly, we offer a
vast collection of books, ranging from classic literary works to
specialized publications, self-development books, and children's
literature. Each book is a new journey of discovery, expanding
knowledge and enriching the soul of the reade
Our website is not just a platform for buying books, but a bridge
connecting readers to the timeless values of culture and wisdom. With
an elegant, user-friendly interface and an intelligent search system,
we are committed to providing a quick and convenient shopping
experience. Additionally, our special promotions and home delivery
services ensure that you save time and fully enjoy the joy of reading.
Let us accompany you on the journey of exploring knowledge and
personal growth!
ebookultra.com