Oxygen and the Evolution of Life

178 Pages·2011·3.556 MB·English

by  Heinz Decker, Kensal E. van Holde (auth.)

Oxygen and the Evolution of Life
.
Heinz Decker Kensal E. van Holde
l
Oxygen and the
Evolution of Life
ProfessorDr.HeinzDecker KensalE.vanHolde
Institutfu¨rMolekulareBiophysik DistinguishedProfessorEmeritus
JohannesGutenberg-Universita¨tMainz DeptofBiochemistryandBiophysics
JakobWelderWeg.26 OregonStateUniversity
55128Mainz,Germany CorvallisOR97331
[email protected] USA
[email protected]
ISBN978-3-642-13178-3 e-ISBN978-3-642-13179-0
DOI10.1007/978-3-642-13179-0
#SpringerHeidelbergDordrechtLondonNewYork
#Springer-VerlagBerlinHeidelberg2011
Thisworkissubjecttocopyright.Allrightsarereserved,whetherthewholeorpartofthematerialis
concerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation,broadcasting,
reproductiononmicrofilmorinanyotherway,andstorageindatabanks.Duplicationofthispublication
orpartsthereofispermittedonlyundertheprovisionsoftheGermanCopyrightLawofSeptember9,
1965,initscurrentversion,andpermissionforusemustalwaysbeobtainedfromSpringer.Violations
areliabletoprosecutionundertheGermanCopyrightLaw.
Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublicationdoesnot
imply, even in the absence of a specific statement, that such names are exempt from the relevant
protectivelawsandregulationsandthereforefreeforgeneraluse.
Cover illustration: Different oxygen transport (respiratory) proteins developed after the oxygen
concentrationincreasedsomebillionyearsago:earthwormhemoglobin(red),arthropodhemocyanin
(scorpion),molluschemocyanin(cephalopod)(frontcover,clockwise)andthemyriapodhemocyanin
(back cover); see also Fig. 5.8. The molecules artwork are courtesy of Ju¨rgen Markl, Institute for
Zoology,JohannesGutenbergUniversityMainz.
Coverdesign:WMXDesignGmbH,Heidelberg,Germany
Printedonacid-freepaper
SpringerispartofSpringerScienceþBusinessMedia(www.springer.com)
Preface
This book has a curious history. It evolved, like its subject, from a much simpler
beginning. Both the authors have had long-standing common interests in the
proteins and processes of oxygen transport in animals. During a sabbatical year
that KvH spent in the laboratory of HD, our discussions broadened to encompass
the much deeper question as to how oxygen transport, and indeed oxygen utiliza-
tion, were related to the evolution of life. As we considered the geological and
paleontological evidence, it became clear that changes in the earth’s atmosphere
andbiologicalevolutionhavebeen,andcontinuetobe,interrelatedincomplexand
fascinatingways.Furthermore,theserelationshipshaveimportantimplicationsfor
humanhealthandhumanity’sfuture.
Thus, the book grew outward from its original focus on oxygen transport,
sometimes into areas in which we must confess less confidence than we would
like.But,wemustaskthereader’sindulgence,forwefeelthatthefascinationofthe
wholestorysuchthatitisvitaltotrytotellit.
One of us (KvH) wishes to express his thanks to the Alexander von Humboldt
Foundation, whose generous support allowed the sabbatical in the Decker labora-
tory. Later, both started the book at the stimulating environment of the Marine
BiologicalLaboratoryatWoodsHolewhereHDspenthissabbatical.
SomereadersmayfindChapter1daunting,withtoomuchdrychemistry.Skipit
ifyouwish!Althoughwefeelthatitprovidesausefulbackgroundfortherestofthe
book,mostofthefollowingChapterscanbereadintelligentlywithoutthismaterial.
We would like to thank Dr. Helmut Ko¨nig, Dr. Wolfgang Mu¨ller-Klieser, and
Dr.HaraldPaulsen(UniversityofMainz)forcriticalreadingofseveralpartsofthe
bookandChristianLozanoskyforhishelpwiththefigures.WealsothankDr.Jutta
Lindenborn(Springer)forallherhelpwiththepublishingprocess.
We would like to express our thanks to our wives, Ina Decker and (the late)
BarbaravanHoldefortheirpatienceduringthepastyears.
Mainz,Germany HeinzDecker
Corvallis,OR,USA KensalE.vanHolde
v
.
Contents
1 Oxygen,ItsNatureandChemistry:WhatIssoSpecialAbout
ThisElement? .............................................................. 1
1.1 ABriefIntroductiontoOxygen ....................................... 1
1.2 AtomicStructureofOxygen:ChemicalBondingPotential .......... 2
1.3 TheDioxygenMolecule ............................................... 5
1.4 ReactiveOxygenSpecies ............................................. 8
1.4.1 Superoxide1O (cid:2)* ............................................... 8
2
1.4.2 Hydrogenperoxide(H O ) ..................................... 9
2 2
1.4.3 Peroxylradical(ROO*) ......................................... 9
1.5 Ozone ................................................................. 10
1.6 Water ................................................................. 12
1.7 WaterVaporintheAtmosphere ..................................... 15
1.8 CarbonDioxide ....................................................... 15
1.9 SolubilityofGasesinWater ......................................... 16
1.10 HydrolysisandDehydration:CentralWaterReactions
inBiology ........................................................... 16
1.11 RedoxReactions ...................................................... 17
References .................................................................. 18
2 ABriefHistoryofOxygen ................................................ 21
2.1 CosmicHistoryoftheElements ....................................... 21
2.1.1 TheSunandSolarSystem ...................................... 24
2.2 FormationofEarth ..................................................... 25
2.3 ThePrimordialEnvironment .......................................... 27
2.3.1 AtmosphereoftheEarlyEarth ................................. 27
2.3.2 WaterontheEarth’Surface:TheOriginofOceans ........... 29
2.3.3 TheFirstGreenhouseEffect .................................... 29
2.4 Life:ItsOriginsandEarliestDevelopment ........................... 30
2.5 ABillionYearsofLifeWithoutDioxygen:AnaerobicMetabolism ... 32
2.5.1 SomePrinciplesofMetabolism ................................ 32
2.6 TheInventionofPhotosynthesis ...................................... 35
vii
viii Contents
2.7 HowOxygenicPhotosynthesisRemodeledtheEarth ................. 38
2.7.1 TheFirstRiseofDioxygen ..................................... 38
2.7.2 EffectsonLife:AnEcologicalCatastrophe? .................. 39
2.7.3 EffectsontheEarth ............................................. 40
References .................................................................. 41
3 CopingwithOxygen ...................................................... 43
3.1 TheImpactofOxygenationonanAnaerobicWorld ................. 43
3.2 ProductionofReactiveOxygenSpecies .............................. 44
3.3 CopingwithReactiveOxygenSpecies ................................ 47
3.3.1 ScavengerMolecules ........................................... 47
3.3.2 EnzymesforDetoxificationofROS ............................ 49
3.3.3 AntioxidantEnzymeSystems .................................. 51
3.4 HowtoAvoidReactiveOxygenSpecies? ............................ 52
3.5 EvolvingDefenseStrategies ........................................... 53
3.5.1 AggregationforDefense ........................................ 53
3.5.2 Melanin ......................................................... 54
3.5.3 OxygenTransportProteinsPreventCreation
ofOxygenRadicals ............................................. 55
3.6 ReactiveOxygenSpeciesasCellularSignals ......................... 56
3.7 DioxygenasaSignal:OxygenSensor ................................ 56
3.8 Summary:ReactiveOxygenSpeciesandLife ........................ 57
References .................................................................. 58
4 AerobicMetabolism:BenefitsfromanOxygenatedWorld ........... 61
4.1 TheAdvantagetoBeingAerobic ..................................... 61
4.2 EvolutionofanAerobicMetabolism .................................. 62
4.2.1 SpecialMechanismsNeededforAerobicMetabolism ........ 62
4.2.2 WhenandHowDidAerobesArise? ........................... 63
4.3 Eukaryotes:TheNextStepinEvolution .............................. 67
4.3.1 DistinctionBetweenProkaryotesandEukaryotes ............. 67
4.3.2 TheSymbioticHypothesis ...................................... 67
4.4 TheLastGreatLeap:MulticellularOrganisms,“Metazoans” ........ 69
4.4.1 When,Why,andHow? ......................................... 69
4.4.2 CollagenandCholesterin ....................................... 70
4.4.3 HalfaBillionYearsofStasis? ................................. 71
4.4.4 EmergenceandExtinctionoftheEdiacaranFauna ............ 72
4.4.5 TheBilateralBodyPlan ........................................ 73
4.4.6 The“CambrianExplosion”:FactorArtifact? ................. 74
References .................................................................. 76
5 FacilitatedOxygenTransport ............................................ 79
5.1 HowtoDeliverDioxygentoAnimalTissues? ...................... 79
5.2 ModesofDelivery .................................................... 80
Contents ix
5.2.1 DiffusionfromtheSurface .................................... 80
5.2.2 TransportviaBloodasaDissolvedGas ...................... 81
5.2.3 OxygenTransportProteins:WhatTheyMustDo? ........... 82
5.3 ModesofDioxygenBindingtoOxygenTransportProteins ........ 84
5.3.1 CooperativeandNoncooperativeBinding .................... 84
5.3.2 HowDoesCooperativityWork?:ModelsforAllostery ...... 86
5.3.3 Self-AssemblyandNesting .................................... 88
5.3.4 WhyComplexMultisubunitOxygenTransportProteins? .... 89
5.4 ModulationofDioxygenDeliverybyOxygenTransportProteins:
Heteroallostery ....................................................... 89
5.4.1 ModulationbytheProductsofAnaerobicMetabolism:
theBohrEffect ................................................. 90
5.4.2 TheHaldaneEffect ............................................ 90
5.4.3 TheRootEffect ................................................ 91
5.4.4 TemperatureDependence ...................................... 92
5.4.5 EvolutionaryAspectsofRegulation ........................... 93
5.5 DiversityofOxygenTransportProteins ............................. 93
5.5.1 Hemoglobins ................................................... 94
5.5.2 Hemerythrins ................................................... 96
5.5.3 Hemocyanins ................................................... 96
5.6 EvolutionofOxygenTransportProteins ............................. 99
5.7 WasSnowballEarthaPossibleTriggerforOPTEvolution? ...... 101
5.8 FromWhatProteinsDidOxygenTransportProteinsEvolve? ..... 102
5.9 OxygenTransportProteinsand“IntelligentDesign” ............... 103
References ................................................................. 103
6 ClimateOvertheAges;IstheEnvironmentStable? .................. 107
6.1 ClimateandGlaciationsinEarth’sHistory .......................... 108
6.1.1 TheFirstMassiveGlaciations;theHuronionEvent:ARole
forMethane? ................................................... 108
6.1.2 LaterProterozoicGlaciations .................................. 110
6.1.3 PhanerozoicClimateandGlaciations ......................... 111
6.2 HowDidLifeSurviveGlaciations? .................................. 116
6.3 MilestonesofLifeinthePhanerozoic ................................ 118
6.4 InorganicCyclingofCarbonDioxide ................................ 121
6.5 IsOurEnvironmentStable? .......................................... 122
6.6 RecentGlobalWarming .............................................. 124
References ................................................................. 124
7 GlobalWarming:HumanInterventioninWorldClimate ........... 127
7.1 RecentClimateChanges .............................................. 127
7.2 PhysicalConsequencesofGlobalWarming ......................... 129
7.2.1 ShrinkingIceandGlaciers .................................... 129
7.2.2 SeaLevelChanges ............................................. 130
7.2.3 ChangesinOceanCurrents .................................... 131
x Contents
7.2.4 LocalClimateandWeather ................................... 132
7.2.5 TheDangerofMethaneReleases ............................. 133
7.2.6 GreenhousetoIcehouseandViceVersa? ..................... 133
7.3 HumanConsequencesofGlobalWarming ........................... 134
7.3.1 DirectConsequencesofCO andTemperatureIncrease ..... 134
2
7.3.2 SeaLevelRise ................................................. 135
7.3.3 ExtremeWeather .............................................. 136
7.3.4 EffectsonAgriculture ......................................... 137
7.4 ControlofGlobalWarming .......................................... 138
7.4.1 PositiveandNegativeNaturalFeedbackMechanism ......... 138
7.4.2 HumanEffectstoControlGlobalWarming .................. 139
7.4.3 TheLongView ................................................ 139
References ................................................................. 140
8 OxygeninMedicine ...................................................... 143
8.1 Hypoxia ............................................................... 143
8.1.1 High-AltitudeHypoxia ........................................ 144
8.1.2 HypoxiaArisingfromMedicalConditions ................... 145
8.2 OxidativeStress ....................................................... 145
8.2.1 NatureofOxidativeStress ..................................... 145
8.2.2 SpecialExamplesofMedicalConsequences
ofOxidativeStress ............................................. 146
8.3 TreatmentofOxidativeStress ........................................ 149
8.4 BeneficialRolesofROS .............................................. 150
8.4.1 SCNandPrimaryImmuneResponse ......................... 150
8.4.2 NitricOxide .................................................... 151
References ................................................................. 153
9 OxygenandtheExplorationoftheUniverse .......................... 157
9.1 WhatIsEssentialfortheDevelopmentofLifeasWeKnowIt? .... 157
9.2 WhatMakesO NecessaryforComplexLifeonHabitable
2
Planets? ............................................................... 158
9.3 SeekingEvidenceforExtraterrestrialLife ........................... 158
9.4 LifeintheSolarSystem? ............................................. 161
9.4.1 TerrestrialPlanets .............................................. 161
9.4.2 IcyMoons ...................................................... 163
9.5 OxygenSupplyProblemsinExtraterrestrialVoyages ............... 164
9.6 ProblemsFacingExtendedExtraterrestrialSettlement
orColonizaton ........................................................ 166
9.6.1 AdjustingthePlanetaryEnvironment:Terraforming ......... 166
9.6.2 AdjustingtheOrganism:Bioforming ......................... 167
References ................................................................. 168
Index .......................................................................... 169