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ARTICLE IN PRESS

Progress in Biophysics and Molecular Biology 89 (2005) 1–8
www.elsevier.com/locate/pbiomolbio

Review

Hypothesis: the origin of life in a hydrogel environment
Jack T. Trevorsa,Ã, Gerald H. Pollackb,1
a

Laboratory of Microbial Technology, Department of Environmental Biology, Ontario Agricultural College, University of Guelph, Guelph, Ontario, Canada N1G 2W1 b Department of Bioengineering, Box 357962, University of Washington, Seattle, WA 98195, USA Available online 22 September 2004

Abstract A hypothesis is proposed that the first cell(s) on the Earth assembled in a hydrogel environment. Gel environments are capable of retaining water, oily hydrocarbons, solutes, and gas bubbles, and are capable of carrying out many functions, even in the absence of a membrane. Thus, the gel-like environment may have conferred distinct advantages for the assembly of the first cell(s). r 2004 Elsevier Ltd. All rights reserved.
Keywords: Assembly; Bacteria; Cell; Clays; Cytoplasm; Division; Earth; Evolution; Growth; Hydrogel; Life; Membrane; Origin; Water

Contents 1. 2. 3. 4. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 The hydrogel environment: superior to a liquid environment for the origin of life? . . . . . . . . . . . . . . . . . . . . 2 Gels and work production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Metabolism in gels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

ÃCorresponding author. Tel.: +1-519-824-4120 ext. 53367; fax: +1-519-837-0442.
1

E-mail addresses: jtrevors@uoguelph.ca (J.T. Trevors), ghp@u.washington.edu (G.H. Pollack). Also to be contacted for correspondence.

0079-6107/$ - see front matter r 2004 Elsevier Ltd. All rights reserved. doi:10.1016/j.pbiomolbio.2004.07.003

. . . . . . . 1992. 1999. . . . . ions. . . . 2001a. . . . . . . . . Cairns-Smith. 2002. . . . . . Nilson. . . . . . . . . . . . . . . deep ocean vents. . Pollack / Progress in Biophysics and Molecular Biology 89 (2005) 1–8 Transition from pre-cell to cell . . as we elaborate below. were ancient life’s most common single-celled forms (Schopf.T. 5 Conclusions. 2002. . . . . . . 1992. . . . . A textbook definition is that the elastic modulus is greater than the viscous modulus: unlike liquids. . Wachtershauser. . . . . . . . . . 7 References . . 1999. . deep hot oily subsurfaces (Gold. . 2004. . We propose that a primitive hydrogel was a more suitable environment for the assembly of precells. . . . . The pre-cell. . . . 2000. . Ingber. . . b. . . Gels. . . . . . . proteins and nucleic acids could remain ordered and in continuous. to controlling drug release in time-release pharmaceuticals. . 6 Acknowledgements . . . . . nutrients. 6. . . . . 2. . .ARTICLE IN PRESS 2 5. many basic functions performed by the cell or pre-cell are capable of being carried out by gels themselves. need not be an issue with the cytoplasm viewed as a cohesive gel. . 2001. . 2001). . Further. . . . . . and for immobilizing enzymes and cells. Trevors and Psenner 2001. . . . . . G. . . ´ 1999)? For what biological. . . . . . . . . . 1999. . . . . . . . 2003a–c. b. . . . Gold. . . . . . . . . . . . . Introduction The origin and evolution of life have been long debated. . . Bashkin. . 1999. . . . . . . J. . . for example. . or in available extraterrestrial samples? In searching for answers to these questions. Morchio and Traverso. . 7 1. . Deamer. . . . . . . . . . . . an implicit presumption is often made that the cell is an aqueous suspension of solutes. . the question of how the pre-cell with no membrane and a very small mass could retain its integrity. . . . . . . retain their integrity even in the absence of a membrane. . . . . . . Morita. as it would in a strictly aqueous environment with free diffusion. . . Trevors. . What. gels do not flow as a result of steady shear. . . . . The hydrogel environment: superior to a liquid environment for the origin of life? Although gels are employed for numerous functions from absorbing urine in disposable diapers. . subsurface cores. or physical life signatures ought one to be searching (Bebie and Schoonen. . . . . . . there is no agreed-upon definition of a gel. . . . . a question worth considering is whether the various conundrums that have limited progress in understanding the origin of life and the genetic instruction set might be better resolved. 2002a. .H. . . . . . Primitive organelles. 2001. Des Marais and Walter. . . Trevors and Abel. Trevors. eventually surrounded by a membrane to become a cell. . . . . . . . . . . and then cells capable of growth and division. . . 1992. . . . . it is known that the cell is a gel (Pollack. . . . . . 2001. .. . With the cytoplasm treated as a gel rather than as an aqueous suspension. . . . . . . . . . . . . . with fundamental questions still unresolved. . . . 1997)? Where should one look for ancient life ¨ ¨ remnants—in thermal springs. then. . . . 1997. . . . A phenomenological definition is that gels are solid or solid-like materials that . . . . is also considered to be an aqueous suspension. chemical. . However. . Hence. . for example. . . . 2001) where water and oil form interfaces. . close molecular physical proximity within the gel without the danger of dissipating. . Segre et al. . . . . . 1985. .

one state is highly hydrated. as it is used in diverse intracellular organelles today (Verdugo et al. Effectively. with a large hydrated diameter of 5 A.. When enmeshed in a polymer network. These changes of volume (or length) are capable of performing work. without a continuous supply of energy. 1994. 1994). one of which is a liquid of some abundance (Almdal et al. Kellermayer et al. Thus. without a membrane. Exclusion is largely size based. the cytoplasmic gel matrix itself is sufficient. as in many common colloidal gels such as gelatin (Bella et al. established the required ion-concentration gradients. Such work could have been used by the pre-cell for metabolism and other critical processes. This property may have provided a mechanism by which the pre-cell. Wiggins. Indeed.ARTICLE IN PRESS J. whereas potassium. Postulated by Dusek and Patterson (1968). such layered. Cameron et al. 1992. water molecules will adsorb onto hydrophilic surfaces. For the pre-cell or primitive cell and even the modern cell. 1999. Gels and work production Unlike the common perception of being largely inert.T. 1965). usage. production. the polymer gel phase transition is a critical phenomenon much like the water–ice transition. the larger the solute the more profoundly the solute will tend to avoid the structured water environment inside the gel/cell (Negendank. 2001). 1993).H. 1984. One feature of hydrogels. it remains unknown if this energy was readily capturable until mechanisms for energy capture. sodium. or ‘‘structured’’ water holds together the network. Similar size-based exclusion of solutes occurs immediately outside cells/gels. discontinuous. Because of both enthalpic and entropic considerations. the solute-exclusion principle has important consequence. One feature of this kind of structured water is that solutes are excluded. Volume changes of 1000 times are not uncommon (Osada and Gong. for distances in the order of 100 mm or more (Zheng and Pollack. Pollack / Progress in Biophysics and Molecular Biology 89 (2005) 1–8 3 consist of two or more components. is that the solvent is not ordinarily bulk water. and amply confirmed by the late Toyo Tanaka (1981) and subsequently by many others. Pollack.. As dipoles. 2001). in which a small change of environment results in a major. gels in water solvent. 1996.. with a smaller hydrated diameter (3. 1993). as for example in artificial muscles (Bar-Cohen. Channels and pumps are not needed. will enter more easily and accumulate. . 1982. and onto one another. The central mechanism of work production is the polymer gel phase transition. A continuous energy supply may or may not have been present when the first pre-cells were being assembled. the other not. 1988. ion partitioning as an equilibrium process had distinct advantages. the hydrogen bonding in water can hold together even polymers that are not cross-linked to one another. forming networks (Ling. Bromberg and Ron. Ling. G. forming a stable gel (Pollack. The principle is commonly exploited in gels known as ion-exchange resins.8 A) and high affinity for protein surfaces (Edelmann. 3. It provides a mechanism for partitioning of critical ions between the ˚ inside and outside. change of physical structure. will be kept from ˚ entering the cell/gel in any quantity.. 2001). Tasaki. Trevors. 2003). 1998). energy is not needed.. the requisite separation is maintained indefinitely. gels are capable of doing work. and storage became available. Hence. 1990).e. i.. If it was. because this is an equilibrium process.

could concentrate solutes. Further. Sun-induced heating could raise the temperature sufficiently to trigger a transition. Thus. all components would be held in a flexible but not too viscous gel that physically maintained components in contact with one and other over long periods of time. shifting the equilibrium towards polymerization and thereby creating conditions more favorable for polymer growth. especially if the enzymes were synthesized slowly and/or were slow acting with the ability to catalyze the transformation of numerous substrates (Demetrius. Pollack / Progress in Biophysics and Molecular Biology 89 (2005) 1–8 Energy for any such transitions would have been derived from the sun. 2001). The mechanism that divides a single bacterial cell into two offspring cells. 1990). microtubules. reactions cannot proceed. Convective flow can be produced by simple propagating phase transitions.H. 4.T. for example. polymer-gel phase transitions offered a simple mechanism by which the pre-cell or early cell(s) could have accomplished various simple tasks. 2001). a gel environment may have provided a more stable environment. Without physical contact. and frequently also by artificial gels that mimic what happens biologically (Pollack. the early presence of a simple mechanism for convective flow would have enhanced the versatility of the pre-cell or early cell. What advantages would a gel-like cytoplasm have offered for these processes. and various types of movement. Phase transitions have been demonstrated to produce various kinds of work relevant to cellular function. 1991). compared to an aqueous cytoplasm? First. Moreover. polymer-gel phase transitions are centrally implicated in basic cellular processes such as secretion. . Central to the assembly of the first cell(s) would have been the mechanisms of energy production. the gel state brought some order to an otherwise disordered environment. potential changes similar to action potentials. A cohesive gel-like cytoplasm is an easier environment to partition into two entities without cytoplasm streaming away. with enzymes remaining active for longer periods of time. this occurs along actin filaments. Thus. to pH and temperature. All of these functions can be accomplished by biopolymer gels. Metabolism in gels The hydrogel environment provided a matrix conducive to non-enzymatic chemical and enzymatic reactions. Enzymatic reactions can occur in narrow. for example. Trevors. and presumably also along similar polymers of a more primitive nature that may have been involved in early cell division (Pollack. while at the same time allowing chemical and eventually enzymatic reactions to occur. Light itself can sometimes trigger phase transitions (Hoffman. either directly or indirectly. nanometer-scale pores. work-producing phase transitions would have been driven by the main source of energy. G. Sun-induced evaporation. Any enzymes necessary for assembly of the first cell(s) that remained active longer would have been more useful than enzymes with a rapid turnover. and also concentrated solutes necessary for the evolution of cellular metabolism. was simpler to encapsulate with an evolving cytoplasmic membrane. in the pre-cell(s). Hence. which could then perform work. The gel environment also facilitates processes such as growth and division.ARTICLE IN PRESS 4 J. 1988). Triggers of the phase transition include any of a vast number of environmental shifts. storage and utilization. where water is almost certainly structured (Wiggins. ranging from ion content. the sun. While the earliest pre-cells may have relied on diffusion alone for nutrient flow. Thus.

5. and also unstable. Even then. Proteins and nucleic acids would still. At some point through an unknown mechanism. molecules. By maintaining a suitable pH. even a simple pentose such as ribose is difficult to make. ions. Gels may have contained various gases useful for metabolic reactions. This liability implies that the first catalytic RNA must have been stabilized in a protective environment. 1995). A hydrogel cytoplasm may also have been conducive to the assembly of the first macromolecules. In addition. Transition from pre-cell to cell The transition between a primitive and a more complex pre-cell may have been the time the cell membrane was first assembled. as well as by hydrolysis reactions. which were likely ubiquitous on the primitive Earth. and rapidly changing temperature. leading to decomposition. osmotic balance and temperature. 1999). It is even possible that the DNA remains more stable. retarding their loss and concentrating them. Gels are also used as a non-restrictive matrix for protein and RNA analysis without evident damage to these macromolecules. such as N2. 1999). while at the same time making them available as substrates in the first cell(s). whereas a strict aqueous solution would have offered less stablity. as are ribonucleosides. In summary. Hydrogen encapsulated in a gel matrix may have been an early substrate for primitive enzymes. eventually oxygen for aerobic respiration) to move in and out without expenditure of energy. the gel environment could have trapped and concentrated gases as they entered the gel. the DNA remaining functional for cloning experiments and polymerase chain reaction (PCR) analysis. degradation can occur in seconds to minutes.T. the gel nature of cells allows diffusion of solutes and gases (e. and coliform bacteria can utilize hydrogen as an energy source but not for growth. however. growth requirements are now fulfilled by other substrates (Morita. The advent of the cell .g. shear forces. pH.ARTICLE IN PRESS J. where stability was challenged by turbulence. H2. Moreover. The ability of DNA to reside as linear DNA or closed circular DNA in present day agarose gels illustrates that a gel environment used in electrophoresis still allows the DNA to take on different configurations. The first gel cytoplasm would have been much simpler as only some dissolved gases. Trevors. ionic concentrations and temperature. Pollack / Progress in Biophysics and Molecular Biology 89 (2005) 1–8 5 accomplishes this amazing activity with no loss of genetic material or cytoplasmic contents. Unless environmental conditions lie within restricted ranges of pH. This type of diffusion is also possible in an aqueous environment.H. CH4 and CO2. The gel matrix may have provided a stable environment for hundreds of millions of years that was not possible in a simple aqueous medium. small peptides. G. increasing their availability for evolving metabolism.. a stable hydrogel cytoplasm may have minimized this difficulty. proteins and catalytic RNA were likely to have been present. once the gases entered the gel environment. the cell surrounded itself by a lipid bilayer. Present day bacteria utilize hydrogen in some reactions. Cytosine is very difficult to synthesize even in laboratory conditions (Shapiro. be capable of folding and taking on their structural conformations in the non-restrictive gel. their diffusion out of the gel will have been slowed. DNA’s stability and functionality are not affected in an adverse manner by the hydrogel environment. thereby defining the cellular entity. however. The hydrogel’s cohesive nature would keep cellular components together. nucleic acids and mRNA are unstable during extraction from cells (Trevors and van Elsas. and chemical concentrations.

channels. In fact. the fraction with the required organelles goes on to divide and produce offspring. The gel holds together. How could the membraneenveloped cell have communicated with the environment in the interim? The gel construct provides a basis to escape from the horns of this dilemma. cell functions need not be seriously affected. Pollack. 6. but also for subsequent cell division and evolution. by what mechanism could the machinery have arisen for the construction of these processes? The pumps and channels were needed instantly.T. within the gel paradigm. as solutes would flow down their respective concentration gradients out of the cell. 1992. the abrupt appearance of the insulating lipid-bilayer membrane presents no conundrum if the bilayer contains orifices such as proteins to facilitate communication. How then could nutrients pass from outside to inside. But development of the manufacturing capability would have taken eons. the cell does not perish (Maniotis and Schliwa. 2001). the gel paradigm offers a solution to the long-held conundrum of how the pre-cell coped before the abrupt appearance of the continuous cell membrane. Despite this violation. The membrane contains a high density of proteins. There is no instant requirement for pumps. We suggest that the . The appearance of the continuous cell membrane presented a largely impermeable barrier between the inside and outside. 1985. but evidently more primitive. One example is that of cells cut into two fractions by the tip of a micropipette. allowing processes to continue.H. by contrast. to avert cell death. than current membrane pumps and channels. Whether continuity is breached by the presence of ordinary leaky proteins. but not for gel structures. Trevors. 1999. The cell would quickly perish. hydrogels have the capacity to carry out various energy-requiring tasks. 1980). Hence. It provided a stable environment not only for the accretion of polymeric mass. Even today’s cells are leaky. 1991). Leaky orifices may present challenges for aqueous suspension models. Tasaki. and there is evidence that phase transitions are used routinely today in highly evolved eukaryotic cells (Verdugo et al. Albrecht-Buehler. routine activity can often continue even though the membrane does not appear to re-grow (Maniotis and Schliwa. the membrane is by no means a pure lipid bilayer. Evidence for this is found in the fact that when additional leakage pathways are created (by even gross membrane violations). Furthermore. 2001). The central point is that the gel holds itself intact whether a continuous membrane is present or not. are not easily tolerated by a cytoplasm that is an aqueous suspension. Such orifices. and waste products in reverse? Accommodating trans-membrane solute flow must have required the presence of membranous components similar to. By undergoing phase transition.. membrane continuity is unnecessary. For vital processes to proceed. it tolerates even large superficial orifices.ARTICLE IN PRESS 6 J. It is riddled with proteins that provide ample leakage pathways. or by manual insult. as much as 80% by weight in bacterial cells and 50% in mammalian cells. This example is but one of a halfdozen classes of ‘‘decapitated’’ cells that nevertheless continue to function (Pollack. The problem is this: with no evolutionary pressure beforehand. or active transport mechanisms. Thus. G. Yawo and Kuno. Such seemingly anomalous behavior is anticipated within the gel construct. Pollack / Progress in Biophysics and Molecular Biology 89 (2005) 1–8 membrane brought a conundrum. Conclusions A hydrogel gel environment provided several advantages for the origin of life. 1991.

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