Phil. Trans. R. Soc.

B (2006) 361, 1845–1856

doi:10.1098/rstb.2006.1908
Published online 7 September 2006

The origin and emergence of life under

impact bombardment
Charles S. Cockell*
Centre for Earth, Planetary, Space and Astronomical Research, Open University,
Milton Keynes MK7 6AA, UK
Craters formed by asteroids and comets offer a number of possibilities as sites for prebiotic chemistry,
and they invite a literal application of Darwin’s ‘warm little pond’. Some of these attributes, such as
prolonged circulation of heated water, are found in deep-ocean hydrothermal vent systems,
previously proposed as sites for prebiotic chemistry. However, impact craters host important
characteristics in a single location, which include the formation of diverse metal sulphides, clays and
zeolites as secondary hydrothermal minerals (which can act as templates or catalysts for prebiotic
syntheses), fracturing of rock during impact (creating a large surface area for reactions), the delivery
of iron in the case of the impact of iron-containing meteorites (which might itself act as a substrate for
prebiotic reactions), diverse impact energies resulting in different rates of hydrothermal cooling and
thus organic syntheses, and the indiscriminate nature of impacts into every available lithology—
generating large numbers of ‘experiments’ in the origin of life. Following the evolution of life, craters
provide cryptoendolithic and chasmoendolithic habitats, particularly in non-sedimentary lithologies,
where limited pore space would otherwise restrict colonization. In impact melt sheets, shattered,
mixed rocks ultimately provided diverse geochemical gradients, which in present-day craters support
the growth of microbial communities.
Keywords: organics; colonization; thermophiles; asteroid; crater

1. INTRODUCTION time (Sleep et al. 1989). Impactors larger than 500 km

During the Hadean, the surface of the Earth was in diameter have been suggested to have sufficient
subjected to relatively heavy bombardment by asteroid energy to boil the whole ocean (Sleep et al. 1989) or to
and comet impacts, compared to the surface of the have at least boiled the top few hundred metres (Ryder
present-day Earth ( Nisbet & Sleep 2001; Kring & 2003). These events would have favoured life in the
Cohen 2002; Ryder 2003). deep regions of the Earth’s crust below the oceans. The
Assuming that the origin of life occurred on Earth, it periodic global heating caused by impacts has been
happened in an environment that was periodically suggested as an explanation for the hyperthermophilic
disturbed. The impact flux on the early Earth was not root of the phylogenetic tree of life (Maher & Stevenson
only higher, but the frequency of large high-energy 1988; Sleep et al. 1989). The hyperthermophilic root
impacts was also higher. Although the impact events neither proves that life originated in hot conditions nor
cause environmental disturbance, the craters associated that the first organism need have been a hypothermo-
with them might have characteristics that make them the phile; the tree of life merely suggests that, at some point
suitable geochemical environments for prebiotic chemi- in early history, a bottleneck resulted in the survival of
stry, particularly with respect to their hydrothermal hyperthermophiles that led to the diversity of life on
systems. The role of impacts in the origin of life has not Earth today (if the root of the phylogenetic tree is a
been examined in any detail. However, hydrothermal reliable indication of the nature of Archaean micro-
vents on the ocean floor have received attention as sites biology, and not merely an artefact of present-day 16S
for the origin of life (e.g. Wächtershäuser 1988; Holm RNA sequences).
1992; Russell & Hall 1997), and these environments, It has been suggested that impact events would have
mutatis mutandis, might provide a basis from which to provided an escape mechanism for life from these early
investigate the potential of impact-induced hydro- ocean-sterilizing impacts. Modelling data suggest that
thermal systems as sites for prebiotic chemistry. micro-organisms could have been ejected into the
Following the emergence of life, impacts would have refugium of interplanetary space on rocks launched
had a profound influence on its distribution and during impact, to return and reseed the Earth several
characteristics. Between the time of the formation of thousand years later ( Wells et al. 2003). Wells’ data
the Earth and ca 3.8 Gyr ago, life in the oceans may suggest that with an initial cell population of 103–105
have been periodically extirpated, if it existed at this cells kgK1, at least one cell in this material would return
after 3000–5000 years, following a sterilizing impact.
Qualitatively similar conclusions have recently been
*[email protected] obtained by Gladman et al. (2005), who show that 1%
One contribution of 19 to a Discussion Meeting Issue ‘Conditions for of the impact-ejected material might eventually return
the emergence of life on the early Earth’. to Earth.

1845 q 2006 The Royal Society

1846 C. S. Cockell Origin and emergence of life

It is apparent, based on the study of extant microbial changes. Thus, craters provide a suite of conditions,
communities within impact-altered target lithologies, which, taken together, make them plausible realistic
that craters would not only have disturbed the early environments for sustained experiments in the origin of
biosphere, but would have provided a suitable, and in life. I will examine each of these characteristics.
some cases improved, geomicrobiological environment
for early life. Certain lithological changes induced by
impact would have improved the conditions for (a) Source of energy and precursors
lithophytic organisms (Cockell et al. 2002). Impactors themselves might have delivered organic
In this paper, I do not propose a chemical theorem precursors for the origin of life (Chyba & Sagan 1992;
for the origin of life, but rather I will examine the Maurette 1998; Charnley et al. 2002), since some
characteristics of impact-induced hydrothermal of these precursors would have survived impact
systems and their associated craters as plausible sites (Pierazzo & Chyba 1999). Organics might have been
for the origin and emergence of life. generated by shock processing of the atmosphere
(Fegley et al. 1986) or during impact itself (McKay &
Borucki 1997), some of which might have been washed
2. CRATERS AND THE ORIGIN OF LIFE into hydrothermal systems from previous impacts, to
Theories on the pathways of prebiotic evolution and be incorporated into the breccias and suevites.
the formation of the first complex self-replicating However, impact hydrothermal systems could have
macromolecules have to take into account several provided conditions for the de novo synthesis of a
common requirements, including: (i) a source of diversity of organic compounds. Numerous previous
energy to drive molecular and macromolecular studies have addressed the possibility of organic
synthesis, (ii) a mechanism for the localized concen- synthesis in deep-ocean hydrothermal systems (e.g.
tration of reactants to favour the required chemical Corliss et al. 1981; Baross & Hoffman 1985; Shock
reactions, (iii) suitable catalysis and (iv) a suitable 1990) and these studies illuminate the potential for
geochemical environment for these reactions and their organic synthesis in impact-induced hydrothermal
products to be sustained for sufficiently long periods to systems. Shock & Schulte (1998), using theoretical
lead to the origin of life. This last requirement is modelling, show that dynamic fluid mixing (as would
important, because although specific chemical have occurred deep in impact craters or following
reactions might be theoretically favoured, these impact events into the Hadean oceans, where hydro-
reactions have to occur in a real environment, not in thermal fluids mixed with Hadean seawater) can yield
laboratories or on a computer. This requires the organic compounds such as carboxylic acids, alcohols
identification of geochemical environments at the and ketones, which are thermodynamically stable in
macroscale that offer promise as sites for the origin of hydrothermal systems. The formation of organic
life at the micro- and nano-scale. compounds from CO2 and H2 within impact hydro-
A diversity of geochemical environments has been thermal systems would have provided a continuous
proposed as suitable sites to meet such requirements. source of precursors for oligomerization reactions, in
Hydrothermal systems have received attention as sites addition to exogenous organics; these latter sources
for organic synthesis and the origin of life (e.g. eventually becoming depleted. The CO2 for hydro-
Wächtershäuser 1988; Ferris 1992; Holm 1992; thermal synthesis would have been derived from the
Russell et al. 1998; Simoneit 2004). At plate Hadean atmosphere and the H2 would have been
boundaries and above magma plumes, hydrothermal
derived from volcanic outgassing and/or the atmos-
fluids would have provided a source of heat and
phere (Tian et al. 2005), or the reaction of water with
chemical energy, in the form of reactive minerals, to
impact-fractured basalts within the crater and its
sustain prebiotic reactions.
fractured surroundings, through the process of serpen-
A sufficient concentration of reactants is difficult
tinization (Moody 1976; Neal & Stanger 1983; Berndt
to imagine in the open oceans, where dilution does
et al. 1996). The impact-induced fracturing of the
not favour the required local conditions for prebiotic
reactions. However, mineral surfaces, such as clays, basement rock could have caused a ‘pulse’ of
can potentially provide templates, surfaces for serpentinization to fuel early organic formation during
sorption, and even catalysis of chemical reactions the onset of the hydrothermal system. Empirical
(Goldschmidt 1952; Rao et al. 1980; Cairns-Smith evidence for the production of hydrocarbons and
1982; Ponnamperuma et al. 1982; Ferris et al. 1988; organics through serpentinization has been presented
Cairns-Smith et al. 1992; Lahav 1994). for the crystalline rocks of the Canadian and
In theories on the origin of life, asteroid and comet Fennoscandian Shields (Sherwood Lollar et al.
impacts generally have been regarded as inimical to 1993) and Conical Seamount in the Mariani forearc
these early prebiotic syntheses. The suggested (Haggerty 1991; Haggerty & Fisher 1992). The
migration of acetogenic precursors of life into the production of hydrocarbons on fresh olivine surfaces
deep subsurface (Russell & Arndt 2005) meets the cracked by volcanically induced expansion and con-
need to escape the hostile surface conditions associated traction has been suggested (Tingle & Hochella 1993),
with impact. indirectly supporting the notion of a role for impact-
However, craters associated with impacts provide induced fracturing in providing surfaces for serpenti-
many required geochemical conditions that favour nization and later hydrocarbon formation, although
prebiotic reactions. Many of these conditions are laboratory experiments simulating the volcanic process
contemporaneous during post-impact environmental have been more equivocal (Tingle & Hochella 1993).

Phil. Trans. R. Soc. B (2006)

 Origin and emergence of life C. S. Cockell 1847

(b) Appropriate minerals for prebiotic reactions reactions. They can catalyse a wide diversity
Impact events are indiscriminate and occur at all of prebiotic reactions (Ferris et al. 1988). The
latitudes and in any lithology that happens to constitute cationic surface of montmorillonite might provide
the target material at the site of the impact. In this template-directed synthesis of RNA oligomers
sense, impacts offer opportunities for many geochem- (Paecht-Horowitz & Eirich 1988; Ferris & Ertem
ical experiments in the origins of life. Although deep- 1992, 1993; Ertem & Ferris 1996).
ocean hydrothermal systems are not homogeneous and A diversity of other hydrothermal minerals found in
different chemical conditions are recognized in craters has been shown to have significance for
different vents (Russell & Arndt 2005), impact craters prebiotic chemistry. In many hydrothermal systems,
potentially offer a larger set of heterogeneous miner- such as those of Popigai crater, Russia; Carswell crater,
alogical conditions for prebiotic reactions, which might Canada and Manson crater, USA, illite is to be found
include early volcanic/granitic lithologies, nascent in the lithic breccias and melt rocks (suevite; Naumov
sedimentary lithologies and mixtures of different 2005). Illite can act as a template for the synthesis of
lithologies, where an impact occurs into a hetero- polypeptides (Hill et al. 1998; Liu & Orgel 1998; Orgel
geneous region or a region with a stratigraphic diversity. 1998). Ferris et al. (1996) synthesized polypeptides
During the period when the origin of life occurred, with over 55 monomers on illite clays.
the diversity of rock types would have been much lower With respect to early Earth, impacts into basalts may
than today; the early crust was probably composed provide the closest analogous geochemical environ-
primarily of komatiites (Nisbet 1987). However, ments, just as organic formation in ultramafic deep-
mobilization of minerals and their subsequent precipi- ocean hydrothermal systems (Holm & Charlou 2001)
tation in the impact hydrothermal system would have are recognized as potentially valuable analogues for
locally enhanced the diversity of secondary minerals early Earth. The 1.8 km diameter, 50 000-year-old
available for prebiotic reactions. Lonar crater, India, resulted from an impact, which
Perhaps, the most significant specific link between occurred into the Deccan traps volcanic province.
impact cratering and reactions for the origin of life are Hagerty & Newsom (2003) examined the hydrothermal
the formation of zeolites and clays as secondary minerals in this crater and found them to be dominated
minerals in impact hydrothermal systems. These by saponite, formed optimally at temperatures between
minerals are formed by aqueous fluids that fill the 130 and 2008C within the post-impact hydrothermal
heated rocks within the crater and react with the shock- system. The ion-exchange properties of saponite
derived aluminosilicates and impact glasses. Clays and minerals are well known (Breen & Moronta 2001) and
zeolites offer charged ordered surfaces on which the potentially offer surfaces for the binding of organic
sorption and oligomerization of reactants can occur, molecules formed within the hydrothermal fluids.
and they have been recognized as important potential These data suggest that within a single impact
templates for prebiotic reactions (Ferris et al. 1988; structure, a great diversity of organic-binding minerals
Zamaraev et al. 1997; Smith 1998; Saladino et al. with different characteristics can be formed, some of
2001). The sorption and ion-exchange properties of which may be more suitable than others as templates.
clays and zeolites make them suitable both as sources The production of other minerals, that are regarded
and sinks for reactants. as suitable surfaces for prebiotic reactions, is favoured
The alteration minerals associated with post-impact in impact craters. Serpentinization can lead to the
hydrothermal systems have now been studied in at least synthesis of double-layered hydroxides, compounds
62 terrestrial impact structures. Altered minerals can which act as ion-exchange surfaces (Cairns-Smith et al.
contribute up to 25% of the mineralogy of the breccia 1992). The pore spaces of weathered feldspar have
formed in a crater. Most of the craters that have been been previously suggested as environments conducive
studied show a great similarity in hydrothermal systems to prebiotic reactions on an account of the ion-
(Naumov 1996), with local variations caused by the exchange properties of this material (Parsons et al.
target lithology. The most common hydrothermal 1998), which might concentrate organics, anions and
minerals to be found are the assemblages of clay cations. The impact-fractured surfaces of feldspar
minerals including smectites with mixtures of zeolites could act as surfaces for prebiotic reactions.
and metal sulphides. On the early Earth, clay minerals Impacts mobilize iron minerals, particularly transition
would have been favoured in the subsurface, where the metal sulphides, with potential consequences for pre-
PCO2 was reduced (Schoonen et al. 2004). biotic reactions (Cody et al. 2001; Cody 2004).
For example, in the Puchezh-Katunki crater, Russia, Hydrothermal systems are known to precipitate metal
deep drilling elucidated the zonation of zeolites with sulphides; these minerals have been found in over 25
Ca/Na and Al/Si ratios increasing downwards into the impact craters (Naumov 2005). The hydrothermal
crater as the temperature of the mineral alterations deposition of vast quantities of economically important
increased (Naumov 1993). Laterally, the zeolites varied metal-rich minerals in large impact structures such as
with zones of high-silica zeolites interspersed with low- Sudbury crater, Canada (iron–nickel ore; Ames et al.
silica varieties. 1998) and Vredefort crater, South Africa (gold ore;
In the upper smectite–zeolite zone of Puchezh- Grieve & Masaitis 1994; table 1) attests to the role
Katunki crater, iron-bearing montmorillonites are to be of impacts in the generation of minerals with significance
found. High Fe-varieties of montmorillonite are found for prebiotic reactions, and the correlation between crater
in the hydrothermal minerals of the Brent crater, size and quantity of minerals deposited. Large impacts on
Canada (Naumov 2005). Montmorillonites have early Earth would have favoured mobilization of metal
received particular attention in studies of prebiotic sulphides. The oxidative formation of pyrite (FeS2) from

Phil. Trans. R. Soc. B (2006)

1848 C. S. Cockell Origin and emergence of life

Table 1. Some characteristics of the impact craters discussed and ion exchange will be rate limiting (Schoonen et al.
in this paper (ages are correct at the time of writing). 2004). The likelihood of reactions occurring will be
greatly improved by an increase in the surface area
diameter available. The impact-induced fracturing of rock vastly
name country (km) age (Myr) increases the surfaces available for reactions. In large
craters, such as Siljan impact structure (52 km
Brent Canada 3.8 396G20 diameter), Sweden, the zone of fractured rock extends
Carswell USA 39 115G10
5 km into the subsurface, and may exceed a volume of
Charlevoix USA 54 342G15
Chesapeake USA 90 35.5G0.3
1000 km3 (Naumov 2005). The fracturing of the rock
Haughton Canada 24 23G1 in impact occurs concomitantly with the establishment
Kärdla Estonia 7 w455 of the post-impact hydrothermal system. In large areas
Lonar India 1.83 0.052G0.006 of the crater, the fresh surfaces are immediately
Manson USA 35 73.8G0.3 exposed to water circulation, and would provide
Popigai Russia 100 35.7G0.2 abundant surfaces for sorption and ion exchange.
Puchezh- Russia 80 167G3 Oligomerization of the precursor molecules into
Katunki more complex molecules, particularly self-replicating
Ries Germany 24 15.1G0.1 ones, is enhanced by non-random attachment of
Siljan Sweden 52 361.0G1.1 molecules to mineral surfaces and it has been shown
Sudbury Canada 250 1850G3
to occur under simulated deep-ocean hydrothermal
Tswaing South Africa 1.13 0.220G0.052
Vredefort South Africa 300 2023G4 conditions ( Imai et al. 1999). The likelihood of these
Wolfe Creek Australia 0.87 !0.3 processes can be increased by a large mineral surface
area on which large numbers of oligomerization
experiments can be accomplished, particularly if the
iron sulphide (FeS) is suggested as a chemoautotrophic reactions are inefficient (Orgel 1998). Large impact
pathway for early life ( Wächtershäuser 1988, 1990). The craters can offer many thousands of cubic kilometres of
mobilization of nickel and other metals in impact craters impact-fracturing rock in contact with hydrothermal
leads to the production of Fe–Ni sulphides and iron oxy- systems for these reactions to occur. As fluids will be
hydroxides, as at Sudbury crater, all of which are known continuously recirculated by convention within an
to promote the reduction of simple precursors of organic impact crater hydrothermal system, the surfaces will
molecules (Huber & Wächtershäuser 1997; Schoonen be refluxed with reactants and oligomerized products,
et al. 1999). leading to further oligomerization on the mineral
Impacts themselves may have delivered metallic Fe surfaces, perhaps much more effectively than in
and Fe–Ni alloys to the surface of the Earth (Schoonen deep-ocean hydrothermal systems, where the reactants
et al. 2004). Meteoritic iron has been shown to enable are continuously being ejected into the dilute sur-
organic syntheses (Gelpi et al. 1970). These com- rounding seawater.
ponents of iron and iron–nickel meteorites would be A disadvantage of impact heated hydrothermal
entrained within the hydrothermal systems and melt- systems is the possibility of high salinities. The briny
sheets of the impact structure created by the impactor water in the deep subsurface of the Chesapeake impact
itself, then the impact event may, in a very real way, structure, USA, has been attributed to hydrothermal
catalyse its own prebiotic reactions. evaporation during the Eocene impact (Sanford 2005).
As high salt concentrations are unfavourable for the
(c) Available surfaces for reactions binding of reactants to mineral surfaces (Orgel 1998),
A part of the kinetic energy of the impactor is manifest regions of impact-induced high salinities, particularly
as shock waves, which alter the target lithology. At high where water becomes trapped and heated within a melt
shock pressures (approx. more than 50 GPa), rock sheet, would be disadvantageous as regions for
melting can occur. Particularly in sedimentary lithol- prebiotic chemistry. However, by contrast, fluid
ogies, these high shock pressures can close pore spaces inclusion data from some craters suggest that the
by both rearranging the grains and melting the hydrothermal fluids of many craters have low (0–14%
material, which flows into the inter-grain space (Kieffer NaCl) salinities (Kirsimaë et al. 2002).
1971; Kieffer et al. 1976). In these cases, we could
expect that surfaces for prebiotic reactions would be (d) Concentrating mechanisms
decreased in abundance and accessibility. However, in The concentration of reactants would occur within
a wide range of lithologies, impact-induced fracturing the hydrothermal system by virtue of deposition and
of the target rock occurs as a result of the propagation sorption on the mineral surfaces with a supply of
of shock waves. In many craters on Earth, an increase in new reactants in hydrothermal fluids and confine-
fracturing in the target lithology has been empirically ment within the fractured rocks as explicated earlier.
observed (e.g. Plado et al. 1996; Pesonen et al. 1999). However, other concentrating mechanisms are
This is usually expressed as a reduction in the density of possible. Near the surface of the crater, evaporative
the rock, an increase in porosity and permeability, and heating and drying of minerals, previously proposed
in some craters, an increase in ground conductivity, as a concentrating mechanism (Usher 1977), would
taken to reflect an increase in saline water within the occur, but in this case induced by hydrothermal
fractured rock. warming. This possibility conforms to the traditional
For a diversity of prebiotic reactions occurring on notion of an evaporative pond as the location for the
the surface of minerals, the number of sites for sorption origin of life, suggested by Charles Darwin (1871).

Phil. Trans. R. Soc. B (2006)

 Origin and emergence of life C. S. Cockell 1849

In the case of continental impacts, the hydrologic and are supersaturated in silica (Naumov 2005). These
depression formed by impact excavation will collect pH values are compatible with several of the prebiotic
water, which subsequently evaporates and provides a reactions described by Wächtershäuser (1990). In
concentrating mechanism. The salty playa in the Wächtershäuser’s scheme, the pH values must be
Wolfe Creek impact crater, Australia, and the highly suitable to allow for high concentrations of HSK,
saline pond in the Tswaing impact crater, South corresponding to pH values greater than 4.5. Many of
Africa (both craters are ca 1 km in diameter), his cleavage reactions are also favoured by non-acidic
provide an empirical demonstration of the ability of pH values. The non-acidic pH is generally more
crater depressions to act as sites for the collection suitable for the surface binding of organics to cationic
and the evaporative concentration of fluids. If the minerals, and the preservation of organic material, for
majority of the Hadean continental crust was any prebiotic reaction sequence. Acidic conditions are
submerged, then these latter evaporative modes of not precluded in impact-induced hydrothermal
concentration will be less important. systems. Osinski et al. (2001) suggest that one
explanation for marcasite deposition in the hydro-
(e) Appropriate physico-chemical conditions thermal systems of the Haughton crater, Canada,
The fracturing, melting and redistribution of target would be reduction of the pH of the hydrothermal
rocks within a crater, both within the faults and within system to below 5. However, geochemical evidence
the melt sheet formed within and around the crater, can suggests that neutral or alkaline pH values are usual for
establish an astonishing diversity of geochemical post-impact hydrothermal systems (Naumov 2005).
gradients and juxtaposed physico-chemical conditions. By contrast, deep-ocean hydrothermal systems
It is recognized, for instance, that different mineral exhibit a diversity of fluid pH values; many of them
types offer particular advantages for the origin of life. are acidic, but some exhibit highly alkaline conditions
Clays provide a surface for attachment and ion (Russell & Arndt 2005), which may themselves inhibit
exchange, and Fe, Co, Cr and Cu-bearing minerals prebiotic reactions.
can provide catalysis for electron-transfer reactions and
organic complexification (e.g. Foustoukos & Seyfried (f ) Longevity of conditions and effects of cooling
2004). These minerals are not necessarily to be found There is no constraint on the length of time required for
in the same location. The fracturing and mixing of the origin of life. An important difference between
rocks prior to their re-emplacement within an impact deep-ocean hydrothermal systems and impact-gener-
structure can bring reactive surfaces into contact with ated hydrothermal systems is the comparatively
one another, generating appropriate, and otherwise temporary nature of the latter. The larger the impact
unusual, physical and chemical conditions over micro- structure, the more long-lived the hydrothermal system
metre scales. will be. For example, at the 24 km diameter Haughton
The hydrothermal system can release catalytic impact structure in Canada, the mineralogical data has
metals from the target rocks, which then interact been interpreted to suggest a hydrothermal system
with templates to catalyse the prebiotic reactions. lasting ca 10 000 years (Osinski et al. 2001). Versh et al.
The hydrothermal release of iron has been demon- (2003) estimated that the hydrothermal system in the
strated in the Charlevoix impact structure, Canada, 4 km diameter Kärdla crater, Estonia, lasted for several
where oxidized iron, together with aluminium thousand years, and Abramov & Kring (2004) suggest
and silicon, is found in the secondary minerals a lifetime of over 2 Myr for the hydrothermal system in
(Trepmann et al. 2005). the 250 km diameter Sudbury impact structure,
These diverse mineral assemblages might improve Canada.
the chances of conditions favourable for oligomeriza- The impact hydrothermal systems start at much
tion of early macromolecules. The minerological higher temperatures than deep-ocean hydrothermal
diversity has been suggested to provide a ‘combina- systems. The initial temperature of the Haughton
torial library’ of systems in which oligomerization could crater hydrothermal system has been estimated at ca
occur (Hill et al. 1998). As different amino acids and 650–7008C (Osinski et al. 2001). Similarly, the Ries
other precursors will adsorb onto different minerals crater (Germany) hydrothermal system may have
with different affinities, depending on charge and local sustained similar temperatures during emplacement
conditions (Liu & Orgel 1998), microenvironments (Pohl et al. 1988). By contrast, on-axis hydrothermal
that offer high geochemical diversity over small systems, which achieve the highest temperatures of
distances will provide environments in which poten- deep-ocean hydrothermal systems, emanate water
tially different and interacting oligomerization temperatures of typically 350–3708C. Thus, impact-
reactions might occur. induced hydrothermal systems will host a potentially
The establishment of thermal gradients within craters, much larger diversity of mineral deposition sequences.
which gives rise to the zonation of secondary minerals, The relatively short duration of impact hydro-
will further lead to a diversity of physico-chemical thermal systems, compared to deep-ocean vents, may
conditions in the active hydrothermal system. We can appear to constrain the opportunity for the origin of
thus think of the disrupted post-impact environment as a life. However, as hydrothermal systems cool, more
‘reactor’ for prebiotic mineral surfaces. complex molecules can form as lower temperatures
Other chemical characteristics of post-impact favour their stability (Shock & Schulte 1998). In
hydrothermal systems lend themselves to prebiotic contrast to deep-ocean hydrothermal vents, the general
reactions. The impact hydrothermal systems typically cooling trend within impact structures creates con-
have a pH of weakly alkaline to near-neutral pH (6–8) ditions, whereby over the lifetime of the hydrothermal

Phil. Trans. R. Soc. B (2006)

1850 C. S. Cockell Origin and emergence of life

system, a large diversity of thermal and cooling life. In the case of impact structures, unlike deep-ocean
regimens are ‘tested’, which potentially corresponds hydrothermal vents, there is no fresh crater existing
to a large diversity of organic syntheses conditions. today in which to directly observe the colonization of
In a diversity of simulated experiments to test organic post-impact hydrothermal systems by extant micro-
synthesis in hydrothermal conditions, too numerous to organisms, but by analogy, we can infer that impact-
adequately review here (e.g. Marshall 1994; Imai et al. induced hydrothermal systems, when cooled to below
1999; McCollom et al. 1999; Ogasawara et al. 2000), it is the upper temperature limit for life, would have offered
apparent that different temperatures favour different environments for early life (Koeberl & Reimold 2004).
syntheses. For example, in simulated laboratory hydro-
thermal experiments run between 100 and 4008C in
stainless steel vessels using oxalic and formic acid as
reactants (as a proxy for H2, CO and CO2), Rushdi & 3. A SCHEME FOR THE ORIGIN OF LIFE WITHIN
Simoneit (2001) showed that alkanoic acids were IMPACT CRATERS
optimally formed at 3008C and alkyl formates at the A tentative scheme for the origin of life within Hadean
lower temperature of 2008C. Other compounds showed impact structures can be proposed. During the
distinctive temperature optima. collision of the asteroid or comet into the oceans or
Theoretical studies also show that different organic exposed land masses, rocks are heavily shocked and
syntheses will be reached at defined temperature fractured in large volumes of the target material. A
optima. The formation of many organic compounds thermal anomaly is generated from the kinetic energy of
in modelled hydrothermal systems is optimal at the object. A hydrothermal system is established within
temperatures of 150–2508C (Shock 1990, 1992) and and around the crater, and secondary mineral depo-
the products of these syntheses, such as acetate, are sition starts to occur. At high temperatures, potentially
generally optimally stable at similar or lower tempera- catalytic metals are released from the target rock and
tures (Mottl 1992). Thus, any geothermal environment deposited within metal pyrites, and zeolites and clays
that allows for temperature variations from several are laid down within the hydrothermal veins and
hundred degrees to much lower temperatures, will fractured rocks.
potentially maximize the diversity of organic syntheses. As the crater begins to cool to temperatures in
As the rate of cooling of impact-induced hydro- the range 50–2508C, several pathways to organic
thermal systems will vary from hours to millions of complexification occur. The presence of CO2 and H2
years, depending on the scale of impact, the target (from the Hadean atmosphere)—H2 being augmented
lithology and the nature of convective fluid flow, this by deep impact-fracturing of Hadean basalts in contact
will further add to the diversity of chemical conditions with seawater, resulting in serpentinization and loca-
tested on Hadean Earth. The optimum conditions for lized formation of H2—would provide the raw material
different reactions will be reached at different times, but for organic synthesis as the hydrothermal fluids come
in the same locations, as the cooling of the hydro- into contact with seawater or fluids within the crater at
thermal system brings the crater through the meta- different temperatures. Alternatively, the reduction of
stable equilibria of aqueous organic compounds. Thus, CO2 could occur in the fluids by catalytic action as it
hydrothermal cooling may even allow for a type of comes into contact with precipitated iron–nickel
‘temporal’ compartmentalization of the reaction hydrothermal minerals in the presence of H2 in the
sequences in the prebiotic period, when cellular scheme proposed by Russell et al. (1998) and
compartmentalization was not available. empirically shown to be capable of producing CH4
By contrast, on-axis deep-ocean hydrothermal under simulated hydrothermal conditions by Horita &
systems have water temperatures much higher than Berndt (1999), or in contact with ferrous sulphide and
those optimum for organic syntheses. As a result, hydrogen sulphide in the schemes proposed by
environments for prebiotic synthesis have focused more Wächtershäuser (1988, 1990). Some of these Fe–Ni
on flank hydrothermal systems (Holm 1992), where reaction sites may be part of iron–nickel impactors
temperatures are more typically ca 1508C. The more themselves and incorporated within the impact breccia
constant temperature regimens would limit the oppor- and suevites. Some organics may already be present in
tunity for diverse reaction sequences in a single the system, generated by shock synthesis during the
location, although temperatures will clearly vary over impact itself and some organics would be present as
spatial scales, but then any mineral-bound reaction material delivered by earlier impactors during this
products will be separated from other reaction products period of heavy bombardment.
produced at different temperatures. Hydrothermal cooling drove the crater environment
through temperature optima for a vast diversity of
(g) Environment for the emergence of life from organic syntheses and complexification reactions. In
the prebiotic reactions some craters, the cooling was too quick for syntheses to
Whatever conditions we postulate for prebiotic produce enough product for subsequent reactions,
reactions, they must be compatible with early life, in whereas in some it was too slow so that reactants from
the sense that the end product of the prebiotic reactions one synthesis were no longer available for later
is a proto-micro-organism that must be capable of syntheses; but in some craters where the kinetic energy
survival, and growth, in the conditions that gave rise to deposited into the target lithology was just right and the
it. Both the deep-ocean hydrothermal vents and the cooling caused by convection and local conditions was
impact-induced hydrothermal systems offer environ- optimal, the organic syntheses of various types would
ments suitable for thermophilic and hyperthermophilic have occurred with optimal rates.

Phil. Trans. R. Soc. B (2006)

 Origin and emergence of life C. S. Cockell 1851

formation of zeolites, clays, metal sulphides and

deposition of other surfaces/catalysts for
prebiotic reactions Hadean
hydrothermal mixing leading Ocean
to organics formation

breccias and suevites

hydrothermal circulation
and cooling

fractured rocks generate H2

fractured and faulted breccia and
basement rocks offering
large surface area for reactions
Figure 1. Darwin’s warm little pond—the impact crater as a prebiotic reactor. Some of the diversity of characteristics of impact
structures that make them favourable sites for prebiotic reactions are shown.

melt rocks and breccias

surface fractured rocks and deposited diverse geochemical gradients for
minerals offer habitats for microorganisms, microorganisms
esp. phototrophs

hydrothermal circulation
for thermophiles in
early stage post-impact
environment

fractured and faulted breccia and

basement rocks offering
large surface area for colonization
Figure 2. Impact craters as a habitat. Some of the characteristics of impact craters that make them favourable sites for micro-
organisms are shown.
The production of organics proceeded to more of one clay/zeolitic reaction being transferred to other
complex stages through the schemes proposed by Russell templates during circulation, resulting in the ‘testing’
and Wächtershäuser and others within the clusters of iron of many oligomerization experiments in diverse phy-
minerals (e.g. pyrite, greigite and mackinawite) laid down sico-chemical conditions, and on diverse substrates
within the hydrothermal systems of the crater. As the throughout the crater.
crater hydrothermal system cooled to temperatures ca Hydrothermal cooling favoured the emergence of
40–1008C, the organic synthesis of acetate, according to more labile complex molecules (Shock & Schulte 1998)
the scheme demonstrated by Huber & Wächtershäuser and at some point within the crater history tempera-
(1997) and also the scheme suggested by Russell, would tures dropped to conditions favourable for complex
have been favourable. nucleic acids, cell membranes and other proto-cellular
Equally plausible, at least from the point of view of structures (e.g. Tsukahara et al. 2002) required for the
the available geochemical environment, is that organics emergence of self-replicating systems.
generated from the various mechanisms available, and Finally, and more speculatively, it is worth noting
described earlier, bound to the impact-generated that these same prebiotic processes may have occurred
zeolites and clays within the crater. During crater in Martian impact craters. Limited tectonic activity on
cooling, oligomerization reactions were favoured early Mars may have significantly constrained the
during the hydrothermal circulation, with the products possibility for deep-ocean type hydrothermal systems.

Phil. Trans. R. Soc. B (2006)

1852 C. S. Cockell Origin and emergence of life

Impact-induced hydrothermal systems may have been

the only means to have generated sustained hydro-
thermal systems for prebiotic reactions.
A schematic of the processes occurring in a crater is
shown in figure 1.

4. CRATERS AS HABITATS FOR EARLY LIFE

Asteroid and comet impact craters offer some
advantages to lithophytic organisms (Cockell et al.
2005; figure 2), and on early Earth, these environ-
ments would have provided suitable places for the
proliferation and radiation of life (Cockell 2004)
once it did emerge. The fracturing of rock during
impact will greatly increase the surface area available
for microbial colonization, although nutrient limi-
tation could limit the degree to which such void
space can actually be used (Cockell et al. 2005),
particularly in the subsurface.
During the early post-impact phase, the crater offers
an environment for what we have previously termed the
‘phase of thermal biology’, defined as that phase
‘during which the thermal anomaly associated with a
recently formed crater sustains biological activity of a
nature or at a level requiring warmer environmental
conditions than would be available in the same area had
the impact event not occurred’ (sensu Cockell & Lee
2002). As adumbrated by Naumov (2005), in most
post-impact hydrothermal environments, these warmer
conditions would be accompanied by neutral to
alkaline pH values, favouring neutrophilic and alkali-
nophilic organisms, although localized regions of acid
conditions are not implausible. The metabolic charac-
teristics of these communities might be analogous to
those that inhabit deep-ocean hydrothermal systems
today (e.g. Nercessian et al. 2003; Alain et al. 2004). As
there is no fresh crater available for study, the true
nature of the hydrothermal microbial ecosystems that
inhabited early Archaean craters must be subject to
some conjecture, but if the earliest life forms were Figure 3. Organisms associated with impact melt sheets
(suevites) in the Haughton impact structure, Canada. (a) A
hyperthermophilic, perhaps owing to an evolutionary
biofilm of organisms associated with a cavity inside the
bottleneck, then impacts would not only have caused material (scale bar 0.1 mm); (b) microfungi associated with
this bottleneck, but also would have generated hot the biofilms (scale bar 30 mm); (c) the diverse mineralogy of
conditions that favoured colonization by the micro- the breccia (granites, dolomite, gneiss, sandstones) provides a
organisms that survived the previous impacts. geochemically diverse environment for microorganisms (scale
The impact-shocked crystalline rocks can provide bar 0.3 mm)
cryptoendolithic environments for diverse organisms.
We have previously shown how, by both increasing
porosity and light transmission, impacts can generate Micro-organisms will also colonize suevites and lithic
novel habitats for photosynthetic micro-organisms breccias. Impact events shatter and mix target rocks.
(Cockell et al. 2002). Habitats within rocks for Some are thrown from the crater, but a considerable
phototrophs are otherwise limited to porous sedimen- volume of fall-back breccias and suevites are deposited
tary lithologies or macroscopic cracks in rocks within the excavation cavity itself. Upon cooling to
(chasmoendolithic communities) on the present-day temperatures favourable for life, the resulting material
Earth. The presence of photosynthetic organisms on contains diverse geochemical interfaces between litholo-
the early Archaean Earth is somewhat equivocal, but gies that may not otherwise be in contact with each
impact-shocked rocks will equally provide a refugium other. In the Haughton impact structure, Canada, we
for heterotrophs and chemolithotrophs, perhaps aiding have observed the presence of filamentous micro-
escape from a putatively harsh UV radiation regimen organisms (presumptive fungi) at the interface between
(Cockell 2004; Cockell & Lim 2005). The minerals laid mudrock emplaced within the suevite and the silicate–
down by impact hydrothermal systems can provide carbonate melt that comprises the matrix (C. Cockell,
sites for microbial colonization. Microbial colonization J. Watson, unpublished data; figure 3). The material
of selenite within the Haughton impact structure has has sufficient porosity to allow for the movement of
been demonstrated (Parnell et al. 2004). micro-organisms into the rock matrix.

Phil. Trans. R. Soc. B (2006)

 Origin and emergence of life C. S. Cockell 1853

On early Earth, mixing of lithologies in this way would Berndt, M. E., Allen, D. W. & Seyfried, W. E. 1996 Reduction
have distributed minerals and established geochemical of CO2 during serpentinization of olivine at 3008C and
gradients (albeit with the probably more limited 500 bar. Geology 24, 351–354. (doi:10.1130/0091-7613
mineralogy available on early Earth compared to (1996)024!0351:ROCDSOO2.3.CO;2)
Breen, C. & Moronta, A. J. 2001 Influence of exchange cation
present-day Earth), which would have provided diverse
and layer charge on the isomerisation of a-pinene over
redox couples for micro-organisms to conserve energy for SWy-2, SAz-1 and Sap-Ca. Clay Miner. 36, 467–472.
growth. Further, geochemical gradients would be (doi:10.1180/0009855013640001)
established between the mixed rocks and the secondary Cairns-Smith, A. G. 1982 Genetic take-over and the mineral
minerals deposited on their surfaces during hydro- origins of life. Cambridge, UK: Cambridge University
thermal circulation in the heavily fractured rocks. Press.
Cairns-Smith, A. G., Hall, A. J. & Russell, M. J. 1992 Mineral
theories of the origin of life and an iron-sulphide example.
5. CONCLUSION Orig. Life Evol. Biosph. 22, 161–180. (doi:10.1007/
Favourable conditions for the origin of life require that a BF01808023)
diversity of factors come together in a single location. Charnley, S. B., Rodgers, S. D., Kuan, Y. J. & Huang, H. C.
Asteroid and comet impact craters offer favourable 2002 Biomolecules in the interstellar medium and in
conditions for prebiotic reactions that are an amalgam comets. Adv. Space Res. 30, 1419–1431. (doi:10.1016/
of conditions, which have previously been postulated as S0273-1177(02)00499-4)
required for the origin of life. The origin of life in the post- Chyba, C. & Sagan, C. 1992 Endogenous production,
exogenous delivery and impact-shock synthesis of organic
impact environment is not an alternative to existing
molecules: an inventory for the origins of life. Nature 355,
theories, but rather offers an expansion of the possible 125–132. (doi:10.1038/355125a0)
number of favourable environments in which such Cockell, C. S. 2004 Impact-shocked rocks—insights into
experiments may have occurred. The environments Archean and extraterrestrial microbial habitats (and sites
described in this paper apply to both hydrothermal for prebiotic chemistry?). Adv. Space Res. 33, 1231–1235.
systems on early cratons and those established on the (doi:10.1016/j.asr.2003.06.027)
seabed following impact into the Hadean oceans. The Cockell, C. S. & Lee, P. 2002 The biology of impact
chemical and the physical conditions, which arise in and craters—a review. Biol. Rev. 77, 279–310. (doi:10.1017/
around impact craters, suggest that Charles Darwin’s S146479310100584X)
‘warm little pond’ (Darwin 1871) has literal relevance in Cockell, C. S. & Lim, D. S. S. 2005 Impact craters, water and
microbial life. In Water and life on Mars (ed. T. Tokano),
understanding possible environments for the origin of
pp. 261–275. Heidelberg, Germany: Springer.
life, and not merely the metaphorical meaning to which it Cockell, C. S., Lee, P., Osinski, G., Horneck, H. & Broady, P.
has, in recent times, been relegated. 2002 Impact-induced microbial endolithic habitats.
After the origin of life, up to the present day, impact Meteorit. Planet. Sci. 37, 1287–1298.
craters offer favourable environments for colonization Cockell, C. S., Lee, P., Broady, P., Lim, D. S. S., Osinski, J.,
by lithophytic organisms. Craters offer a localized Parnell, J., Koeberl, C., Pesonen, L. & Salminen, J. 2005
source of liquid water, mobilization of minerals and Effects of asteroid and comet impacts on habitats for
carbon and, in melt sheets and suevite, a potentially lithophytic organisms—a synthesis. Meteorit. Planet. Sci.
large diversity of geochemical gradients, particularly 40, 1901–1916.
where the lithologic target sequence is diverse, Cody, G. D. 2004 Transition metal sulfides and the origins of
subaerial and submerged craters would have offered metabolism. Ann. Rev. Earth Planet. Sci. 32, 569–599.
(doi:10.1146/annurev.earth.32.101802.120225)
favourable environments for diverse microbial com-
Cody, G. D., Boctor, N. Z., Hazen, R. M., Brandes, J. A.,
munities on early Earth. Morowitz, H. J. & Yoder, H. S. 2001 Geochemical roots of
I acknowledge the NASA Haughton-Mars Project, under autotrophic carbon fixation: hydrothermal experiments in
whose auspices the microbiological work described here was the system citric acid, H2O–(GFeS)–(GNiS). Geochim.
conducted. Cosmochim. Acta 65, 3557–3576. (doi:10.1016/S0016-
7037(01)00674-3)
Corliss, J. B., Baross, J. A. & Hoffman, S. E. 1981 An
REFERENCES hypothesis concerning the relationship between submar-
Abramov, O. & Kring, D. A. 2004 Impact-induced hydro- ine hot springs and the origin of life on Earth. Oceanol.
thermal system at the Sudbury crater: duration, temperatures, Acta 4, 59–69. Proc. 26th Int. Geol. Congress, Geology of
mechanics, and biological implications (abs). Lunar and the Oceans symposium, Paris, France.
planetary science XXXV. Houston, TX: LPI no. 1697 Darwin, C. 1871 Letter to Hooker, J. D.
(CD-ROM). Ertem, G. & Ferris, J. P. 1996 Synthesis of RNA oligomers on
Alain, K., Zbinden, M., Le Bris, N., Lesongeur, F., heterogeneous templates. Nature 379, 238–240. (doi:10.
Querellou, J., Gaill, F. & Cambon-Bonavita, M. A. 2004 1038/379238a0)
Early steps in microbial colonization processes at deep-sea Fegley, B., Prinn, R. G., Hartman, H. & Watkins, G. H. 1986
hydrothermal vents. Environ. Microbiol. 6, 227–241. Chemical effects of large impacts on the Earth’s primitive
(doi:10.1111/j.1462-2920.2003.00557.x) atmosphere. Nature 319, 305–308. (doi:10.1038/319305a0)
Ames, D. E., Watkinson, D. H. & Parrish, R. R. 1998 Dating of a Ferris, J. P. 1992 Marine hydrothermal systems and the origin
regional hydrothermal system induced by the 1850 Ma of life: chemical markers of prebiotic chemistry in
Sudbury impact event. Geology 26, 447–450. (doi:10.1130/ hydrothermal systems. Orig. Life Evol. Biosph. 22,
0091-7613(1998)026!0447:DOARHSO2.3.CO;2) 109–315. (doi:10.1007/BF01808020)
Baross, J. A. & Hoffman, S. E. 1985 Submarine hydrothermal Ferris, J. P. & Ertem, G. 1992 Oligeromerisation reactions
vents and associated gradient environments as sites for the of ribonucleotides on montmorillonite—reaction of the
origin and evolution of life. Orig. Life Evol. Biosph. 15, 5 0 -phosphorimidazolide of adenosine. Science 257,
327–345. (doi:10.1007/BF01808177) 1387–1389.

Phil. Trans. R. Soc. B (2006)

1854 C. S. Cockell Origin and emergence of life

Ferris, J. P. & Ertem, G. 1993 Montomorillonite catalysis of Kieffer, S. W., Phakey, P. P. & Christie, J. M. 1976 Shock
RNA oligomer formation in aqueous solution: a model for processes in porous quartzite: transmission electron
the prebiotic formation of RNA. J. Am. Chem. Soc. 115, microscope observations and theory. Contrib. Mineral.
12 270–12 275. (doi:10.1021/ja00079a006) Petrol. 59, 41–93. (doi:10.1007/BF00375110)
Ferris, J. P., Huang, V.-H. & Hagan, W. J. 1988 Montmor- Kirsimaë, K., Suuroja, S., Kirs, J., Kärki, A., Polikarpus, M.,
illonite: a multifunctional mineral catalyst for the pre- Puura, V. & Suuroja, K. 2002 Hornblende alteration and
biological formation of phosphate esters. Orig. Life Evol. fluid inclusions in Kärdla impact crater, Estonia—an
Biosph. 18, 121–133. (doi:10.1007/BF01808786) indication for the post-impact hydrothermal activities.
Ferris, J. P., Hill, A. R., Liu, R. H. & Orgel, L. E. 1996 Meteorit. Planet. Sci. 37, 449–457.
Synthesis of long prebiotic oligomers on mineral surfaces. Koeberl, C. & Reimold, W. U. 2004 Post-impact hydro-
Nature 381, 59–61. (doi:10.1038/381059a0) thermal activity in meteorite impact craters and potential
Foustoukos, D. I. & Seyfried, W. E. 2004 Hydrocarbons in opportunities for life. IAU Symp. 213, 299–304.
hydrothermal vent fluids: the role of chromium-bearing Kring, D. A. & Cohen, B. A. 2002 Cataclysmic bombard-
catalysts. Science 304, 1002–1005. (doi:10.1126/science. ment throughout the inner solar system 3.9–4.0 Ga.
1096033) J. Geophys. Res. 107. (doi:10.1029/2001JE001529)
Gelpi, E., Han, L., Nooner, D. W. & Oró, H. 1970 Closed- Lahav, N. 1994 Minerals and the origin of life—hypotheses
system Fischer–Tropsch synthesis over meteoritic iron, and experiments in heterogeneous chemistry. Heterogen.
iron ore and nickel–iron alloy. Geochim. Cosmochim. Acta Chem. Rev. 1, 159–179.
34, 965–979. (doi:10.1016/0016-7037(70)90079-7) Liu, R. & Orgel, L. 1998 Polymerisation on the rocks:
Gladman, B., Dones, L., Levison, H. F. & Burns, J. A. b-amino acids and arginine. Orig. Life Evol. Biosph. 28,
2005 Impact seeding and reseeding in the inner Solar 245–257. (doi:10.1023/A:1006576213220)
System. Astrobiology 5, 483–496. (doi:10.1089/ast. Maher, K. A. & Stevenson, D. J. 1988 Impact frustration of
2005.5.483) the origin of life. Nature 331, 612–614. (doi:10.1038/
Goldschmidt, V. M. 1952 Geochemical aspects of the origin 331612a0)
of complex organic molecules on the Earth, as precursors Marshall, W. L. 1994 Hydrothermal synthesis of amino acids.
to organic life. New Biol. 12, 97–105. Geochim. Cosmochim. Acta 58, 2099–2106. (doi:10.1016/
Grieve, R. A. F. & Masaitis, V. L. 1994 The economic 0016-7037(94)90288-7)
potential of terrestrial impact craters. Int. Geol. Rev. 36, Maurette, M. 1998 Carbonaceous micrometeorites and the
105–151. origin of life. Orig. Life Evol. Biosph. 28, 385–412. (doi:10.
Hagerty, J. J. & Newsom, H. E. 2003 Hydrothermal 1023/A:1006589819844)
McCollom, T. M., Ritter, G. & Simoneit, B. R. T. 1999 Lipid
alteration at the Lonar lake impact structure, India:
synthesis under hydrothermal conditions by Fischer–
implications for impact cratering on Mars. Meteorit.
Tropsch-type reactions. Orig. Life Evol. Biosph. 29,
Planet. Sci. 38, 365–381.
153–166. (doi:10.1023/A:1006592502746)
Haggerty, J. A. 1991 Evidence from fluid seeps atop
McKay, C. P. & Borucki, W. J. 1997 Organic synthesis in
serpentine seamounts in the Mariani Forearc: clues for
experimental impact shocks. Science 276, 390–392.
the emplacement of the seamounts and their relationship
(doi:10.1126/science.276.5311.390)
to forearc tectonics. Mar. Geol. 102, 293–309. (doi:10.
Moody, J. B. 1976 Serpentinization—a review. Lithos 9,
1016/0025-3227(91)90013-T)
125–138. (doi:10.1016/0024-4937(76)90030-X)
Haggerty, J. A. & Fisher, J. B. 1992 Short-chain organic acids
Mottl, M. J. 1992 Pore waters from serpentinite seamounts in
in interstitial waters from Mariana and Bonin forearc
the Mariana and IzuBonin forearcs, Leg 125: evidence for
serpentinites: Leg 125. Proc. ODP, Sci. Results 125, volatiles from the subducting slab. Proc. ODP, Sci. Results
387–395. 125, 373–385.
Hill, A. R., Böhler, C. & Orgel, L. 1998 Polymerisation on Naumov, M. V. 1993 Zeolite mineralization in impact carters.
the rocks: negatively charged a-amino acids. Orig. Life Zapiski Vsesoyuznogo Mineralogicheskogo Obshchestva 122,
Evol. Biosph. 28, 235–243. (doi:10.1023/A:100657 1–12.
2112311) Naumov, M. V. 1996 Basic regularities of the post-impact
Holm, N. G. 1992 Why are hydrothermal systems proposed hydrothermal process. Sol. Syst. Res. 30, 21–27.
as plausible environments for the origin of life? Orig. Life Naumov, M. V. 2005 Principal features of impact-generated
Evol. Biosph. 22, 5–14. (doi:10.1007/BF01808015) hydrothermal circulation systems: mineralogical and
Holm, N. G. & Charlou, J. L. 2001 Initial indications of geochemical evidence. Geofluids 5, 165–184. (doi:10.
abiotic formation of hydrocarbons in the Rainbow 1111/j.1468-8123.2005.00092.x)
ultramafic hydrothermal system, Mid-Atlantic Ridge. Neal, C. & Stanger, G. 1983 Hydrogen generation from
Earth Planet. Sci. Lett. 191, 1–8. (doi:10.1016/S0012- mantle source rocks in Oman. Earth Planet. Sci. Lett. 66,
821X(01)00397-1) 315–320. (doi:10.1016/0012-821X(83)90144-9)
Horita, J. & Berndt, M. E. 1999 Abiogenic methane Nercessian, O., Reysenbach, A. L., Prieur, D. & Jeanthon, C.
formation and isotope fractionation under hydrothermal 2003 Archaeal diversity associated with in situ samplers
conditions. Science 285, 1055–1057. (doi:10.1126/science. deployed on hydrothermal vents on the East Pacific Rise
285.5430.1055) (13 degrees N). Environ. Microbiol. 5, 492–502. (doi:10.
Huber, C. & Wächtershäuser, G. 1997 Activated acetic acid 1046/j.1462-2920.2003.00437.x)
by carbon fixation on (Fe,Ni)S under primordial con- Nisbet, E. G. 1987 The Young Earth—introduction to Archean
ditions. Science 285, 245–247. (doi:10.1126/science.276. geology. Boston, MA: Allen and Unwin.
5310.245) Nisbet, E. G. & Sleep, N. H. 2001 The habitat and nature of
Imai, E.-I., Honda, H., Hatori, K., Brack, A. & Matsuno, K. early life. Nature 409, 1083–1091. (doi:10.1038/350
1999 Elongation of oligopeptides in a simulated sub- 59210)
marine hydrothermal system. Science 283, 831–833. Ogasawara, H., Yoshida, A., Imai, E.-I., Honda, H., Hatori,
(doi:10.1126/science.283.5403.831) K. & Matsuno, K. 2000 Synthesizing oligomers from
Kieffer, S. W. 1971 Shock metamorphism of the Coconino monomeric nucleotides in simulated hydrothermal
sandstone at Meteor crater, Arizona. J. Geophys. Res. 76, environments. Orig. Life Evol. Biosph. 30, 519–526.
5449–5473. (doi:10.1023/A:1026539708173)

Phil. Trans. R. Soc. B (2006)

 Origin and emergence of life C. S. Cockell 1855

Orgel, L. E. 1998 Polymerization on the rocks: theoretical and amino acids with the FeS/FeS2 redox couple as
introduction. Orig. Life Evol. Biosph. 28, 227–234. (doi:10. reductant. Orig. Life Evol. Biosph. 29, 5–32. (doi:10.1023/
1023/A:1006595411403) A:1006558802113)
Osinski, G. R., Spray, J. & Lee, P. 2001 Impact-induced Schoonen, M., Smirnov, A. & Cohn, C. 2004 A perspective
hydrothermal activity within the Haughton impact on the role of minerals in prebiotic synthesis. Ambio 33,
structure: generation of a transient, warm, wet oasis. 539–551.
Meteorit. Planet. Sci. 36, 731–745. Sherwood Lollar, B., Frape, S. K., Weise, S. M., Fritz, P.,
Paecht-Horowitz, M. & Eirich, F. R. 1988 The poly- Macko, S. A. & Welhan, J. A. 1993 Abiotic methanogen-
merisation of amino acid adenylates on sodium-montmor- esis in crystalline rocks. Geochim. Cosmochim. Acta 57,
illonite with pre-adsorbed peptides. Orig. Life Evol. Biosph. 5087–5097. (doi:10.1016/0016-7037(93)90610-9)
18, 359–387. (doi:10.1007/BF01808216) Shock, E. L. 1990 Geochemical constraints on the origin of
Parnell, J., Lee, P., Cockell, C. S. & Osinski, G. R. 2004 organic compounds in hydrothermal systems. Orig. Life
Microbial colonization in impact-generated hydrothermal Evol. Biosph. 20, 331–367. (doi:10.1007/BF01808115)
sulphate deposits, Haughton impact structure, and Shock, E. L. 1992 Chemical environments of submarine
implications for sulphates on Mars. Int. J. Astrobiol. 3, hydrothermal systems. Orig. Life Evol. Biosph. 22, 67–107.
247–256. (doi:10.1017/S1473550404001995)
(doi:10.1007/BF01808019)
Parsons, I., Lee, M. R. & Smith, J. V. 1998 Biochemical
Shock, E. L. & Schulte, M. D. 1998 Organic synthesis during
evolution II: origin of life in tubular microstructures on
fluid mixing in hydrothermal systems. J. Geophys. Res. 103,
weathered feldspar surfaces. Proc. Natl Acad. Sci. USA 95,
28 513–28 527. (doi:10.1029/98JE02142)
15 173–15 176. (doi:10.1073/pnas.95.26.15173)
Simoneit, B. R. T. 2004 Prebiotic organic synthesis under
Pesonen, L. J., Elo, S., Lehtinen, M., Jokinen, T., Puranen, R.
& Kivekäs, L. 1999 Lake Karikkoselkä impact structure, hydrothermal conditions: an overview. Adv. Space Res. 33,
central Finland: new geophysical and petrographic results. 88–94. (doi:10.1016/j.asr.2003.05.006)
In Large meteorite impacts and planetary evolution II (ed. Sleep, N. H., Zahnle, K. J., Kasting, J. F. & Morowitz, H. J.
B. O. Dressler & V. L. Sharpton), pp. 131–147. Boulder, 1989 Annihilation of ecosystems by large asteroid impacts
CO: Geological Society of America Special Paper. 339. on the early Earth. Nature 342, 139–142. (doi:10.1038/
Pierazzo, E. & Chyba, C. F. 1999 Amino acid survival in large 342139a0)
cometary impacts. Meteorit. Planet. Sci. 34, 909–918. Smith, J. V. 1998 Biochemical evolution. I. Polymerisation on
Plado, J., Pesonen, L. J., Elo, S., Puura, V. & Suuroja, K. internal, organophilic silica surfaces of dealuminated
1996 Geophysical research on the Kärdla impact zeolites and feldspars. Proc. Natl Acad. Sci. USA 95,
structure, Hiiumaa Island, Estonia. Meteorit. Planet. Sci. 3370–3375. (doi:10.1073/pnas.95.7.3370)
31, 289–298. Tian, F., Toon, O. B., Pavlov, A. A. & De Sterck, H. 2005
Pohl, J., Eckstaller, A. & Roberston, P. B. 1988 The Ries A hydrogen-rich early Earth atmosphere. Science 308,
impact crater. In Impact and explosion cratering (ed. D. J. 1014–1017. (doi:10.1126/science.1106983)
Roddy & R. O. Pepin), pp. 343–404. New York, NY: Tingle, T. N. & Hochella, M. F. 1993 Formation of reduced
Permagon Press. carbonaceous matter in basalts and xenoliths: reaction of
Ponnamperuma, C., Shimoyama, A. & Friebele, E. 1982 C–O–H gas on olivine cracked surfaces. Geochim.
Clay and the origin of life. Orig. Life Evol. Biosph. 12, 9–40. Cosmochim. Acta 57, 3245–3249. (doi:10.1016/0016-
(doi:10.1007/BF00926908) 7037(93)90537-7)
Rao, M., Odom, D. G. & Oró, J. 1980 Clays in prebiological Trepmann, C. A., Gotte, T. & Spray, J. G. 2005 Impact-
evolution. J. Mol. Evol. 15, 317–331. (doi:10.1007/ related Ca-metasomatism in crystalline target-rocks from
BF01733138) the Charlevoix structure, Quebec, Canada. Can. Mineral.
Rushdi, A. I. & Simoneit, B. R. T. 2001 Lipid formation by 43, 553–567.
aqueous Fischer–Tropsch-type synthesis over a tempera- Tsukahara, H., Imai, E.-I., Honda, H., Hatori, H. &
ture range of 100–4008C. Orig. Life Evol. Biosph. 31, Matsuno, K. 2002 Prebiotic oligomerisation on or inside
103–118. (doi:10.1023/A:1006702503954) lipid vesicles in hydrothermal environments. Orig. Life
Russell, M. J. & Arndt, N. T. 2005 Geodynamic and Evol. Biosph. 32, 13–21. (doi:10.1023/A:1013962328048)
metabolic cycles in the Hadean. Biogeosciences 2, 97–111. Usher, D. A. 1977 Early chemical evolution of nucleic acids: a
Russell, M. J. & Hall, A. J. 1997 The emergence of life from theoretical model. Science 196, 311–313.
iron monosulphide bubbles at a submarine hydrothermal Versh, E., Kirsimäe, K, Jöeleht, F., Plado, J. 2003. Impact-
redox and pH front. J. Geol. Soc. Lond. 154, 377–402.
induced hydrothermal system at Kärdla crater—
Russell, M. J., Daia, D. E. & Hall, A. J. 1998 The emergence
development and biological consequences (abs). Third
of life from FeS bubbles at alkaline hot springs in an acid
International Conference on Large Meteorite Impacts.
ocean. In Thermophiles: the keys to the molecular evolution
Nordlingen, 5–7 August, 2003, no 4040 (CD-ROM).
and the origin of life (ed. J. Wiegel & M. W. W. Adams),
Wächtershäuser, G. 1988 Pyrite formation, the first energy
pp. 77–126. Washington, DC: Taylor & Francis.
Ryder, G. 2003 Bombardment of the Hadean Earth: source of life: a hypothesis. Syst. Appl. Microbiol. 10,
wholesome or deleterious. Astrobiology 3, 3–6. (doi:10. 207–210.
1089/153110703321632390) Wächtershäuser, G. 1990 Evolution of the first metabolic
Saladino, R., Crestini, C., Costanzo, G., Negri, R. & Di cycles. Proc. Natl Acad. Sci. USA 87, 200–204. (doi:10.
Mauro, E. 2001 A possible prebiotic synthesis of 1073/pnas.87.1.200)
purine, adenine, cytosine, and 4(3H)-pyrimidinone from Wells, L. E., Armstrong, J. C. & Gonzalez, G. 2003
formamide: implications for the origin of life. Bioorg. Reseeding of early Earth by impacts of returning ejecta
Med. Chem. 9, 1249–1253. (doi:10.1016/S0968-0896(00) during the late heavy bombardment. Icarus 162, 38–46.
00340-0) (doi:10.1016/S0019-1035(02)00077-5)
Sanford, W. E. 2005 A simulation of the hydrothermal Zamaraev, K. I., Romannikov, V. N., Salganik, R. I.,
response to the Chesapeake Bay bolide impact. Geofluids Wlassoff, W. A. & Khramtsov, V. V. 1997 Modelling of
5, 185–201. (doi:10.1111/j.1468-8123.2005.00110.x) the prebiotic synthesis of oligopeptides: silicate catalysts
Schoonen, M., Xu, Y. & Bebie, J. 1999 Energetics and help to overcome the critical stage. Orig. Life Evol. Biosph.
kinetics of the prebiotic synthesis of simple organic acids 27, 325–337. (doi:10.1023/A:1006571800690)

Phil. Trans. R. Soc. B (2006)

1856 C. S. Cockell Origin and emergence of life

Discussion to hydrothermal vents in deep sea and impact craters as

J. I. Lunine (Lunar and Planetary Sciences Department, environments for emergence of life. How do these
University of Arizona, USA). Your excellent treatment environments compare to each other? Does life need
of terrestrial hydrothermal systems can be extended to high temperature only present in hydrothermal vents,
environments on Mars and Saturn’s Moon Titan as but NOT in tidal basin ponds?
well. On Mars, the abundant impact craters in crust C. S. Cockell. Tidal basins have always been a favoured
that holds water ice might create transient liquid water location for the origin of life. We obviously do not know
environments, where prebiotic chemistry could be whether the origin of life needed high temperatures,
hosted perhaps through to the origin of life. On but high temperatures do provide favourable con-
Titan, impacts into the organic bearing water ice ditions for organic syntheses (assuming these were
crust would have provided liquid water for hundreds
required endogenously) and for speeding up organic
or thousands of years, in which the organic molecules
reactions (although not too hot or high temperatures
might have undergone interesting prebiotic reactions.
will break down organic compounds). Tidal pools offer
There’s a beautiful 450 km diameter crater captured by
the advantage of possible evaporative concentration of
Cassini radar on Titan that shows evidence for
reactants on mineral surfaces or just in solution,
reworking, channels and possibly infill or flooding. It
although one can imagine concentration of reactants
is a candidate for a return mission to Titan.
in hydrothermal veins as they become attached to
This is correct that impact hydrothermal systems
may be places for complex organic chemistry on other mineral surfaces. To evade your question—yes, each
planets and indeed there has been some work in this environment does have some advantages and disad-
direction already. Of course, if the hydrothermal vantages, but I think we just do not know enough about
systems produce complex organic compounds, then the origin of life to be able to say ‘this environment is
they are very interesting, but much less interesting if the much more likely than another’.
opportunity is not there for the emergence of life F. Diego. Why is the temperature decay after the
following the complexification of the organics. impact so slow; i.e. from 500 to 1008C in as long as
C. S. Cockell. I should say that in the context of 10 000 years?
applying these ideas to other planets, it is the case that C. S. Cockell. The decay of temperature obviously
owing to the lack of, or the short duration of, plate depends on the scale of the impact. Remember that
tectonics on Mars, or indeed, any Mars-like planet the kinetic energy squares with the velocity and these
elsewhere, where deep-ocean hydrothermal systems at objects are coming in at over 10 km sK1. That is a lot
spreading centres may not exist, then impact-induced of energy delivered into the bed rock. The rate of
hydrothermal systems may be even more important as cooling will be influenced by convection, the local
sites for these early reactions, maybe even the only sites. climate and many other factors. However, that time
I focused my discussion on Earth owing to the focus of period of 10 000 years is estimated for a crater of
the discussion meeting, but your comments on the about 25 km across. Larger craters such as Sudbury,
relevance of these discussions to other planets are very which is over 200 km across, would potentially drive
valuable. hydrothermal systems for over a million years. It
F. Diego (Department of Physics and Astronomy, seems slow, but that’s quite quick geologically and the
University College London, London, UK ). Can you question is, of course, is such a time period too quick
comment on the conditions in tidal basins compared to be useful for the origin of life?

Phil. Trans. R. Soc. B (2006)