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Home , Cell death, Hierarchy of death, Immortality, Programmed cell death

Christian Spirituality and Issues in the Contemporary World

Volume 9 Issue 1 Subjective Reality, Complexity and Article 3 Programmed

2013

Genetically Programmed : Concepts of Death and in the of the

Paul U. Cameron Monash University, [email protected]

Lynden Rogers Avondale College of Higher Education, [email protected]

Follow this and additional works at: https://research.avondale.edu.au/css

Recommended Citation Cameron, P. U., & Rogers, L. (2013). Genetically : Concepts of death and immortality in the age of the genome. Christian Spirituality and Science, 9(1), 32-55. Retrieved from https://research.avondale.edu.au/css/vol9/iss1/3

This Article is brought to you for free and open access by the Avondale Centre for Interdisciplinary Studies in Science at ResearchOnline@Avondale. It has been accepted for inclusion in Christian Spirituality and Science by an authorized editor of ResearchOnline@Avondale. For more information, please contact [email protected]. Cameron and Rogers: Genetically Programmed Cell Death: Concepts of Death and Immortal

Genetically Programmed Cell Death: Concepts of Death and Immortality in the Age of the Genome. Associate Professor Paul U Cameron, Clinical Immunologist and Immunopathologist Infectious Department, Monash University Department, Alfred Hospital Infectious Diseases Unit, Alfred Hospital Melbourne, Victoria

and

Lynden J Rogers School of Science and Mathematics Avondale College of Higher Education Cooranbong, NSW

ABSTRACT Scientific discoveries have done much to complicate matters of and death. Perhaps the most significant such revelation in recent decades has been that every cell in our bodies has a built-in, and genetically controlled capability. This is also true of most other . Cells know how to die! It has been discovered that programmed cell death turns out to be vitally important in many life processes such as: the development of the mature from a single cell, combatting foreign threats, such as , and controlling rampant cell growth, as found in various . It is also important in the normal of skin and gut lining. Cell death appears to be much more inextricably intertwined with life than may have been earlier supposed. This paper introduces this discussion and explores some of the implications of programmed cell death for a Christian understanding of the relationship between life and death.

Keywords: , programmed cell death, , immunity

1. INTRODUCTION death and sorrow are no more, there One of the dominant motifs woven is an underlying concept of death as through the Judeo-Chri­ stian world an enemy from which can view is that of death as the ultimate be saved.3 Indeed, one of the best expression of evil, a foe that must and known verses in the canon, John 3:16, will be overcome.1 From the Genesis encapsulates this view.4 Furthermore, account of the fall and the expulsion many Christians understand not just from Eden2 through to the Revela- death but death in any form and tion account of a new world where under all circumstances, to be a result

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of sin’s “curse”. However, it may be by the realisation that the outer layers asked how well this view of death as of skin, for example, are comprised of the ultimate enemy accords with the dead cells. observations of modern biology. All this has changed. Biological 2. ISSUES OF LIFE AND science has revealed that all living DEATH organisms, both and , are There is considerable evidence to sug- composed of small basic units called gest that the idea of death in biblical cells. These work in similar ways in was very simple and was associ- plants and animals and are fundamen- ated only with the world of animals and tal to all processes of life and death for humans. Animals were either “quick” both. We also now recognise the exist- (moving) or “dead”, as in Acts 10:425. ence of a huge of . Certainly, life and death were holistic Some of these are beneficial, even in that these states applied to the whole essential, to higher life forms while organism at once. Texts such as Gen others are deadly. Furthermore, we 9:4, Lev 17:13, 14 and Deut 12:236 understand that living entities utilise suggest also that, for the Hebrews, many dead cells, such as those found life was closely associated with blood. in the outer layer of skin, in their quest Indeed, the whole sanctuary ritual re- for survival, and continually produce flected this understanding. This may them. So in this sense even living suggest that only animals with blood things are not completely “alive”. Of were regarded as being alive. Inter- course, science still has much to learn estingly, some evidence suggests that about life and death. Viruses provide plants, being stationary in any case, an illustration of the difficulty of actu- were not thought of as being alive or ally defining life. Although possessing dead in the same way as were animals; a small amount of either DNA or RNA they were either “green” or “dry”.7 contained within a coat, they do Clearly, such an understanding would not have any intrinsic and have allowed the animals and humans replicate only inside the cells of other in the Garden of Eden, all of which ac- organisms. For these reasons they are cording to the literal reading of the text not regarded as actually being “alive”. were vegetarian, to eat without causing Additionally, Australia has many fasci- death. That such ingestion and diges- nating life forms, such as cryptobiotic tion may have involved the systematic midge larvae, which can lie dormant elimination of microorganisms within and desiccated for years under dry the gut represented no problem since outback river beds, to all intents and all such processes were invisible to purposes dead and showing no visible those in Bible times. Similarly, with sign of metabolism. Yet a sudden, no concept of the “cell”, they would soaking rain can “wake” them up, not have been theologically challenged whereupon they quickly become active

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and reproduce, all in the few days the ments or ecological niches, yet each puddles remain.8 Just what differenti- is quite restricted in the range of ates such an organism in a quiescent conditions it can tolerate.10 , state from one which is actually dead although successful in populating is not well understood at all. Such dis- various hostile environments because coveries have enormously complicated of their ability to generate body heat our comprehension of death. and their utilisation of various forms of insulation, have cellular building One breakthrough which has proved blocks which are particularly limited central to an understanding of many in this regard. Isolated mammalian areas of , including repro- cells are very fragile compared to, duction and immunology, concerns for example, , and can only the programmed death of cells. There exist momentarily outside the body. is now an explosion of scientific lit- Also, they die very easily if exposed erature on this topic. A milestone in to low temperatures, as evidenced by this understanding was marked by the the death of the fingers, toes or other awarding of the Nobel Prize in medi- extremities because of frostbite. Ex- cine in 2002 to , Sydney posure to high temperature is similarly Brenner and Robert Horvitz for their damaging. Small children can suffer work on the genetic control of cell severe consequences from high fevers, development and death.9 What exactly while a rise in body temperature of as did these do and why did little as 5 degrees C can produce heat their discoveries merit such recogni- and death in hot, dry, desert tion? What is programmed cell death environments. Many bear the scars and why is it so significant? What is of encounters with hot objects such the relationship between death at the as boiling water, stove hotplates or single cellular level and the death of fires. These result from the violent an organism? Finally, what are the death of individual cells making up implications of the existence of such a the affected tissue. biological system for our reading of the Genesis account, particularly concern- Such cell death results not only from ing the effect of early human on physical damage but also from meta- death throughout the biosphere? This bolic insults such as exposure to carbon paper addresses these questions. First, monoxide (from car exhausts), cya- we need to differentiate between two nide, or the loss of − oxygen, main types of cell death. glucose, essential amino acids, protein components and vitamins. From what- 3. NECROSIS AND ever cause it may arise, this traumatic APOPTOSIS: DIFFERENT form of cell death is called “necrosis” TYPES OF CELL DEATH (Greek necros, meaning “dead body”). Living organisms are found within a relatively broad range of environ- But over the last 30 years

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have become aware of another, more may be seen under a microscope as a deliberate, mechanism of cell death, ruffling of the cell membrane which one quite distinct from necrosis. This proceeds to clumping or condensation is programmed cell death. John Kerr, of the and eventually to the an Australian pathologist whose inter- development of a featureless cell cyto- est was in research, and his plasm with dense fragments of nuclei. associates were the first to coin a term This nuclear fragmentation gave rise designating this process in pathologi- to one of the first instrumental tests cal specimens.11 He suggested the term for identifying the process of apop- “apoptosis” (Greek apoptosis, mean- tosis. The method initially involved ing “falling off”). The simplest way to extracting DNA from a cell thought to demonstrate cell death by this means is be dying by apoptosis and separating to deprive cells in culture medium of it according to size by running it on the blood serum that is normally added an electrophoretic gel in order to al- to maintain cell growth. This was, low the size of DNA segments to be in fact, the way it was first observed determined. If this produced a “lad- experimentally. The process of apop- der” of DNA fragments with multiples tosis is in effect a systematic suicide of 180-200­ bases then apoptosis was of the cell according to a genetically confirmed. More modern methods use pre-programmed path. sophisticated measures that detect the One of the earliest markers of apop- actual ends of the DNA fragments, the tosis is a characteristic change in the number of which is greatly increased nucleus which contains the governing due to the breaking up of the DNA genetic material of the cell, the DNA. during apoptosis. These sensitive The DNA is associated with techniques enable the identification called histones, which produce struc- of apoptosis in individual cells. The tures called nucleosomes. Each nu- nuclear fragmentation is eventually cleosome protects a DNA span of some followed (generally some hours later) 180-200 nucleotide bases, so that there by modifications to the cell surface is a small gap of "naked'' DNA every which allow the apoptotic cells to be 200 or so bases. It was soon discov- recognised by phagocytic cells (cells ered that in apoptosis a DNase enzyme which engulf dead or harmful cells and cuts the unprotected DNA between the organisms), leading to their ingestion nucleosomes, producing a DNA “lad- and complete destruction. der” made up of multiples of these 200 While apoptosis was the first form of or so bases. This DNase activity was programmed cell death to be identified clearly due to some other cell enzyme by microscopy, and has been well- produced by the cell itself and was characterised biochemically, other obviously internally, i.e. genetically, forms of programmed cell death are controlled. This process of apoptosis

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also now known to exist.12 It is now the cell cluster, not only by a process of clearly understood that the mecha- controlled but one of nisms controlling programmed cell regulated cell loss. From the formation death are genetically controlled and of the cavity in the onwards, are “built in” to life forms at a very apoptosis has been found to be critical fundamental level. The award of the in this moulding process. For example, 2002 Nobel Prize to Sulston, Brenner in a developing human embryo, “be- and Horvitz was a recognition of their tween the forty-sixth and fifty second important roles in providing vital in- days in the womb the interdigital web- sights into these processes. bing of the hand suddenly disappears, leaving behind five beautifully shaped It was rapidly appreciated that the fingers.”13 process of apoptosis is vital to normal embryological development and of Another instance that demonstrates critical importance to the immune sys- the fundamental importance of pro- tem. It also proved to be important in grammed cell death in embryology curbing abnormal cell growth, as in the concerns human development. case of cancers. Related processes of During the second trimester there is programmed cell death have also been huge proliferation of neurons within found to be important in regulating the the . About half of normal processes of renewal in many these are subplate neurons, a special tissues, including the skin and gut. category which form some of the Sections 4−7 of this paper describe the first functional cortical brain circuits. role of programmed cell death in these They function in some sense as neural significant life processes. “scaffolding”, and provide crucial regulation of cortical development and 4. PROGRAMMED CELL plasticity. During the third trimester DEATH IN EMBRYOLOGICAL and early post-natal life most of these DEVELOPMENT subplate neurons die by apoptosis, All multicellular organisms develop their essential work done.14 It is pos- from a single cell, a fertilised ovum. sible that a failure of these apoptotic The embryo advances from a single regulatory mechanisms may be as- cell to a cluster of cells (), next sociated with some forms of autism.15 to a ball of cells with a central cavity (blastocyst), then to a plate of cells 5. APOPTOSIS IN THE which eventually develops into the layers of cells (gastrula) that progres- One of the most important survival sively differentiate to form a head and mechanisms for multicellular organ- a tail end. This then folds to produce a isms is the discrimination between . In each of these steps there self, harmless variations of self and must be remodelling of the structure of nonself.­ A further task is to differen-

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tiate non-self organisms which do not encounter with a foreign organism pose any particular threat, such as food or cell and to respond rapidly and components, from those which may effectively on subsequent exposure. threaten survival. The immune system The increased efficiencies of such an is the body's defence barrier that makes immune system have contributed to the these decisions and determines an ap- success of mammals in a wide range of propriate response. environments and in particular to their ability to colonise new environments The development of genetically pro- rapidly where novel parasites and mi- grammed mechanisms for cell death crobes pose a threat. Without adaptive provides a number of “hooks” into immunity, protection from a pathogen which the immune system can link requires changes in the genetic reper- in order to kill foreign, abnormal or toire. Since genome changes can only virus-infected cells. It has been found occur as a result of differential survival that the immune responses in multicel- such genetic shifts are slow and may lular organisms comprise a number require many . On the of tiered layers. Simple organisms other hand, adaptive immunity, with exhibit only the lower, “innate” levels, its of previous exposures to whereas more advanced life forms, harmful organisms, allows survival such as mammals, whilst retaining the and thus the colonisation of a new en- more simple mechanisms, also utilise vironment within a single . a range of “adaptive” mechanisms. These are much more sophisticated It was who recognised and effective. It should be noted at that lymphocytes (small white blood this point that although the term “im- cells) were responsible for this memo- mune” most correctly relates only to ry. In 1960 he and Macfarlane Burnett the consequences of these adaptive received the Nobel Prize for their mechanisms, the terms “immune description of tolerance and immune system” and “immune response” are recognition and their identification frequently, and somewhat loosely, of clonal selection of lymphocytes as used to describe all functions within the basis of immunological memory.16 the body’s defence system. However, (Before their observations were pub- in this brief treatment we consider only lished it was thought that these cells adaptive immunity. Not surprisingly, were somehow “instructed” to respond the complex signalling pathways and to foreign material only after exposure processes found within the human to that material.) However, although immune system have been particularly the existence of immunological memo- well defined. ry and tolerance was established by the 1960s, the basis for such recognition Adaptive immunity involves the was not established until much later. system’s ability to recall a previous

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It was soon realised that these defence Controlled mechanisms could be activated by The human genome contains only organ transplants. In fact, one of the about 20,000 - 25,000 different protein earliest recognised examples of this coding . These genes must code phenomenon was graft rejection. It for embryonic development as well as was noted that if skin from one mouse many details concerning cell and tissue was grafted onto a genetically different function for the life of the organism. mouse it would be rejected, generally Even without these additional (and after a delay of some days. If a second most important!) functions there is no graft from the same donor was applied possibility that such a small number some later it would be rejected of genes could individually code for within minutes or hours. However, if protection against not only pathogens the second graft was from a new donor but each of the almost infinite number the rate of the rejection process was no of possible antigen configurations different from the first graft. Thus the which might pose a threat. So how memory for graft exposure is specific does the immune system prepare itself for each donor. The number of react- to counter all such eventualities? ing cells is increased during exposure The answer to this dilemma is found to a graft and when re­exposed to a in the process of controlled genetic graft from the same source this large recombination, which involves the population of reactive cells recognises generation of lymphocytes with recep- the foreign proteins and rejects them tors which have a wide range of recog- rapidly. nition specificity from which selection The characterisation of the action of can be made. This occurs within two lymphocytes acting in the immune parallel systems. For T lymphocytes response of graft rejection has now led (T cells), it takes place in the thymus, to the recognition of several different a small organ of the immune system subgroups within these lymphocytes, which is located behind the sternum all contributing to a complex interac- near the heart. For B lymphocytes tion that leads to adaptive immunity (B cells) this occurs in the bone mar- and memory. There are three stages row. These cells have similar genetic to the process of protection through coding mechanisms that provide for adaptive immunity: controlled genetic the generation of a huge amount of recombination; clonal selection of variation in surface receptors. These those cells that show reactivity to the include recombination from multiple foreign material; and the subsequent small genes and then the introduction elimination of the threat. Both of the of diversity by a random joining last two stages involve apoptosis. which produces a bewildering array of “fixes”, mostly to problems which will never be encountered! It transpires that

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a small , such as a tadpole, hav- tors. Both killer T cell (cytotoxic T ing as few as one million lymphocytes, cells) and helper T cell responses use can generate an immune response to the same T cell receptors. For B cells a virtually any protein to which it might variant form of the secreted antibody or be exposed, although in an animal of immunoglobulin produced by that cell this size there may only be one cell in is used as a on the cell surface. the whole body that can recognise the When such recognition occurs the foreign material. Thus, are next stage, for both T and B cells, considered to have essentially an un- involves the selective replication of limited repertoire or ability to respond those cells that interact most strongly to foreign proteins. with the foreign protein. Thus this Clonal Selection cell proliferation is highly specific for Some cells generated by this process the foreign protein or antigen. This of genetic modification turn out to expansion of cells takes place by clonal be “self reactive”, i.e. reactive to the selection over 5 to7 days, during which host organism. If they were allowed there may have been 8 to10 cycles of 17 to circulate they could destroy huge doubling . The expanded population numbers of perfectly functional cells, of cells, however, is long-lived so that thus creating havoc. This does in fact when the foreign material is encoun- happen in the case of certain autoim- tered again there are enough cells to mune diseases. Such cells must be respond immediately, thus eliminating quickly eliminated and are, in fact, the threat. In this sense the expanded removed by apoptosis. clonal cell line provides “memory” of that specific antigen and the animal is Once launched on their defensive said to be “immune”. career the remaining randomly spe- cific lymphocytes need the stimulus Elimination of the Threat by of a “recognition” event in order to Apoptosis survive. As earlier noted, for a very The cells killing the organ transplant, large majority of them this never oc- or other outside “threat” as the case curs. After a time all such cells die by may be, do so first by recognising the apoptotic suicide. cells as foreign and then, fast on the draw, delivering a killing signal to However, for some lymphocytes a rec- the foreign cell which initiates pro- ognition event will occur. This recog- grammed cell suicide. The cell under nition is mediated by on the attack has no defence, and will die. surface of the lymphocytes which are the products of the rearranged genes. 6. APOPTOSIS AND CANCERS On T cells these are T cell receptors Cancer is an unregulated expansion and on B cells they are B cell recep- of cells. This is usually initiated by in genes that control cell

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growth, lifespan or death, i.e. so-called name and so are known as “HeLa” “cancer genes”. Normally, damage to a cells.18 Usually a cell will accumulate cell by: typically found in mutations in several such genes before tobacco smoke, UV light, isotopic par- malignancy becomes evident. ticles or gamma radiation consists of Thus cancers can result from changes altered nucleotide bases in one strand in a number of cancer genes, which of the cellular DNA. However, there direct and control apoptosis. Cancer are two such strands and a number of genes can be categorized into sev- mechanisms exist for repairing breaks eral groups: tumour suppressor genes, or modifications in one of these. If genes that produce growth factors, these strategies cannot restore the DNA repair enzymes and genes in- original form then permanent damage volved in apoptotic pathways. Clearly, is caused. These changes to the DNA if there is a loss of these regulatory code may result in the production of mechanisms then any irregular cell abnormal proteins and hence altered growth may proceed unchecked. In cell growth. Cells which accumulate fact, the study of the of cancer too much damage to the DNA may has been one of the main areas show- enter the apoptotic path and politely ing the important role of apoptosis in destroy themselves for the good of the regulation of cell numbers and cell the organism. However, this does not growth and thus of apoptosis as a target always occur. for drug therapy. The genes that are mutated in cancer 7. PROGRAMMED CELL DEATH cells and which result in increased IN NORMAL SKIN AND GUT growth include , a particular As noted earlier, unprotected cells are sub-population of genes that normally physically fragile and if exposed to the regulate cell growth but can be altered, environment will rapidly die. Cells allowing accelerated or uncontrolled need to be kept within a temperature cell growth. Some abnormalities range of about 30 to 40 °C and kept are associated with loss of the normal moist in a solution with nutrients process of apoptotic cell death so that and defined salts. Cells kept in fresh the cells proliferate without death and water will swell by osmosis until they thus become immortal. This was the disintegrate. Conversely, if immersed case for , cells from in sea water they will shrink, also by whose cancerous tumour famously be- osmosis, and die. Some sort of cover- came one of the most heavily used cell ing is needed to protect cells in most lines in the biological research world. multicellular organisms. Crustaceans According to convention these cells have a carapace, a hard external shell are identified by the first two letters to protect the animal from the external of the donor’s given name and family environment, while land molluscs

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produce secretions (slime) that in- In all of these systems cells damaged sulate them from their environment. by environmental insults are lost and Mammals have a system of physical replaced without causing damage to protection based on skin and growths the organism. Problems occur when from this skin (hair or wool). there is significant damage to the proliferating cells in the basal layers. The outermost layer of skin is a beau- For example, genetic forms of bowel tifully crafted system composed of cancers result from abnormalities of dead cells which can cope with the genes that control either the prolif- environmental insults to which they eration or programmed death of these are frequently exposed. These dead gut-lining cells. cells are continually lost by abrasion and must be replaced. They do so by 8. SUMMARY OF CURRENT continual cell division in the bottom UNDERSTANDING OF layer of the skin. These cells then ac- PROGRAMMED CELL DEATH cumulate a substance, , within A number of conclusions can be made. each cell and eventually undergo a These include the following key points. form of sequential, programmed cell • There is a complex system death called “cornification”. In this composed of many genes which situation the complete destruction and program cell death mechanisms. elimination of the cells, as in apoptosis, is not desirable, since they are required • Programmed cell death pathways as a barrier.19 A similar mechanism op- exist in all organisms from yeast erates in the bladder, mouth and throat. to man.

In the gut the mechanism is somewhat • Without programmed cell death similar but there the lining consists of a the normal embryological single layer of cells rather than stacked development of an organism layers. Growth and proliferation oc- from a single cell, the fertilised curs in the crypts, glands located at ovum, into a mature adult could the bottom of and between the villi, not occur. which are the small projections into the • Apoptosis is critically important food flow within the gut. The lining in the development of the cells produced in the crypts gradually immune system. Without it the migrate up to the tips of the villi where adaptive immune response and their numbers are controlled by pro- immunological memory could grammed cell death mechanisms and not exist. abrasive loss into the gut. In this way there is continual renewal of the cells • The removal of cells by lining the gut as they are worn away apoptosis is vital to the by passing food. protection of the organism from

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the development of tumours and of a cell’s death depend on whether cancers that might otherwise or not its death benefits or harms, result in its death. even kills, the organism as a whole? This question subtly undergirds any • Programmed cell death modern discussion of the morality of mechanisms are involved in life and death. the normal development and maintenance of skin and gut One might also ask whether the ces- lining. sation of life processes in a cell can be regarded as being of the same 9. THE RELATIONSHIP moral significance as the death of the BETWEEN CELL DEATH organism as a whole. At a reduction- AND THE DEATH OF THE ist level the answer is probably “yes”. ORGANISM However, at the holistic level, where The life of an organism is a conse- the whole is regarded as more than quence of, and is evidenced by, the simply the sum of the parts, the life continuation of a number of activities of a sophisticated organism may be and processes carried out at the cellular regarded as a higher entity, emergent level. Furthermore, there are a number from continuing cellular processes, but of vital body systems in which the mal- morally differentiable from the “life” function or death of key cells brings possessed by the constituent cells. In about rapid, even instant death of the this case the answer may well be “no”. organism as a whole. In this sense life However, even in the latter case some on Earth, even in its highest human modifications may be required to the dimensions of sentience, conscious- traditional Christian understanding of ness and awareness of the spiritual, is life and death processes operative in ultimately dependent on the successful “the beginning”. Certainly, the way continuity of cellular processes. Of this question is answered might influ- course, while the demise of these key ence the comparative appeal of the cells brings death to the organism other models discussed below. cells can die with little or no ill effect, sometimes even with benefit. In fact, 10. IMPLICATIONS FOR as has been shown, mechanisms of CHRISTIAN THOUGHT programmed cell death contribute for 10.1 The Origin of Programmed the most part to the normal develop- Cell Death Mechanisms ment and survival of the organism The complex genetic system20 control- as a whole. One might ask whether ling programmed cell death comprises there is any moral difference between hundreds of the estimated 20,000 – the programmed death of a useless or 25,000 genes in the human dangerous cell, or the incidental death genome21 and is similarly represented of a vital cell. Does the significance in all lower . These cell death

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processes are present from life’s be- been introduced by God after the fall. ginning to its end. Death at this level It may also be possible to construct appears to be an essential part of life! intermediate models between these Although not the subject of this paper, extremes. We will refer to these as we also know that the same is true at “mixed” models. the level of ecological systems.22 All growth and proliferation is balanced Pre-Fall Model by death and decay. This dependence This model assumes that the original of life on death allows stable systems creation did indeed incorporate all to exist, dependent only on energy the genetic machinery governing input from the Sun. From a purely programmed cell death and that not scientific perspective it is difficult to only embryological development but conceive of life without death. This immune responses and protection raises some obvious theological issues from malignancy occurred in the pre- for Christians. human-fall created order. It would also seem most consistent to posit the pro- Modern Christians face two main cesses of cell death and the consequent possibilities for the origin of the death of larger organisms as being es- programming which controls life and sential aspects of this order. In other death at the cellular level. Either the words, life for most of the creation then sophisticated genetic system currently was very much like life now. Under controlling cell death in all species this scenario mankind would have been was created by divine fiat (creationist the only creature for which immortality model) or it has evolved stepwise in was possible, this being a “higher-level response to the requirements for cell state'' imposed on the natural order by survival during the gradual develop- the “''. Once mankind fell ment of organisms from simple to and was excluded from this tree then complex (evolutionary model). These the natural order governed humanity, contrasting scenarios are explored just as it had all life outside the garden. below, with a particular focus on crea- As far as the authors are aware, only tionist models. one Adventist author has described a model anything like this.23 10.2 Creationist Models For creationists two main possibili- The pre-fall model is clearly successful ties stand out. What might be called in some respects. the “pre-fall” model suggests that all • The proposed picture of cell these mechanisms must have been death and the death of higher present as part of the original creation organisms would enable the before the fall. However, according to existence “in the beginning” the “post-fall” model all mechanisms of stable ecosystems, waste for programmed cell death must have

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recycling and population limits, It also suffers from some significant all of which are very consistent failures. with the findings of modern • Most obviously and importantly science. it does not accord with the • It also provides an obvious role traditional paradigms of for the Tree of Life, namely the universal perfection and eternal inhibition of the natural order of life which many Christians death and recycling for mankind. understand to be associated with Indeed, the most obvious reading both the beginning and end of of Gen 3:22 is that prolonged human history according to a life for humanity was dependent variety of scriptural passages.27 upon eating from the Tree of • It may be difficult to reconcile a Life. This model also makes it God whose original fiat creation easy to understand why access to included such features as it was forbidden after the fall. programmed cell death with the • It may also be understood to be character of Christ, who claimed in harmony with other aspects of to be the “ and the a literal reading of Genesis. The life” incarnate. context of Genesis 1 & 2 can be understood to imply the creation Post-fall Model of a special garden, outside of This model assumes that the initial which is a natural order in need creation was entirely free from any of “subduing”24. Perhaps the fact death or decay. This model is often that, according to the narrative, portrayed in conservative Christian Adam was not created in the literature. According to this view all Garden but later brought into life came forth perfect and immortal, it,25 may also indicate that Adam albeit conditionally, from the Creator’s did not have intrinsic eternal life hands and there could have been no and that he would need a Tree of death or decay at any level of life’s Life. hierarchy before the fall. Accordingly, all programmed cell death mechanisms • It may also be consistent with would have been established by God those biblical statements in in a post-fall creation “recall”, in order which God appears to be to assist the maturation and survival comparatively unconcerned with of organisms now facing the many the death of animals, or with deadly environmental insults brought 26 . about by man’s fall. This “re-creative” • It is entirely consistent with all work must have taken place over a known functions of cell death. short time, perhaps again by divine

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fiat, since there is insufficient time organisms, against which in traditional Christian time scales defence would be required. to allow sequential development. In However, it also suffers from some any case, most Christians taking this shortcomings. view understand the Genesis account to be describing a sudden rather than • Since programmed cell death a gradual transformation of . appears to be inextricably According to this view, mechanisms implicated in the embryological of programmed cell death would be development of all new life, it seen essentially as the curse of man’s is difficult to understand how sin being visited upon creation at large. new generations could have arisen in pre-fall Eden. Yet Although not within any awareness those supporting this model of programmed cell death as such, recognise that God clearly bade historical Adventism could be said to his “perfect” creation be fruitful. have identified most strongly with this model. Despite the dissemination of • Further, it appears somewhat knowledge concerning programmed ironic that mechanisms of cell death over recent decades the programmed cell death should writers are unaware of any serious Ad- have been employed in the ventist attempt to integrate these data promulgation of life: the into the “all death post-fall” traditional embryological development of view. This model has been assumed new life and with the protection rather than defended by most modern of that life from environmental Adventist writers on Origins. insult.

This view is again successful in many • Another problem arises from the respects. perception of some adherents that pre-fall flowers may have • Since the cause of all death is been everlasting. Ellen White identified as Satan, sin and the wrote that the Earth as it came , God’s character is from the Creator’s hand “bore no exonerated from any first-cause blight of decay or shadow of the responsibility for death in any curse”.28 Also suggestive of the form. same sentiment is her description • Clearly, too, this view is of the scene immediately post- compatible with at least the fall: “As they witnessed in immunological function of drooping flower and falling leaf programmed cell death since the the first signs of decay, Adam “fallen” world could be expected and his companion mourned . . to contain many threatening . The death of the frail, delicate

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flowers was indeed a cause for have simply been a visible sorrow; but when the goodly metaphor of eternal life, as it is trees cast off their leaves, the presumably in the New Earth.31 scene brought vividly to However, such an interpretation the stern fact that death is the goes somewhat against a literal portion of every living thing.”29 reading of the text. Since seeds today cannot • Pre-fall life must have operated form or germinate without the according to fundamentally programmed death of the cells different microbiological comprising the flower it is principles and mechanisms difficult to understand how pre- and therefore the post-fall fall occurred modifications required to build at all. If it did, it must have been in and manage mortality must by mechanisms very different have been fundamental and from those currently operating. extensive. Yet changes on such • This view also struggles to a scale are not obvious from the understand the function of description of this process in the Tree of Life. The context scripture. suggests that the Tree of Life existed only for humans, Mixed Models since no mention is made of It is clearly possible to construct its intended consumption by hybrids of the Post-fall and Pre-fall animals. Why make it available models described above. For exam- just to humans? Further, if ple, programmed cell death processes both animals and humans may have been part of the original were intrinsically free of death creation but were intended simply to processes what purpose was assist in embryological development, served by having this tree at good digestion and insulation from the all? Some have conjectured environment, benign and congenial as that humans alone may have it was. Without any embarrassment been created without intrinsic such a composite model might also rec- immortality and that this ognise as inevitable the programmed would have been “awarded” death of plant cells and the drooping of by God after a suitable period flowers in the process of fruit and grain of obedience,30 whereupon the maturation. Such instances of cell tree may have been redundant death would not be seen as a result or and would, perhaps, have manifestation of evil but simply as the been withdrawn. The Bible, sensible and sustainable way in which however, is silent on this point. God chose to build His “perfect” and Perhaps, too, the Tree may fully functional world. Such models

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would draw a clear distinction between this waste – presumably by the likes of death at the cellular level and the death bacteria and dung beetles doing pretty of a higher organism, particularly a much what they do today. He sug- conscious, sentient, death-aware or- gested that, according to such a view, ganism such as a human being! White’s comments concerning the post-fall decay might refer principally According to these mixed models, to the “loss of strength, soundness, programmed cell death mechanisms health and beauty” rather than to the might have been adapted by God death of plant cells, including those after the fall and utilised to meet the of flowers.32 emergent threats, such as pathogens and runaway cell proliferation, which Such composite models achieve some were somehow induced by man’s fall. resolutions.

Although not specifically discussing • They still identify the death programmed cell death, Leonard Brand of higher organisms as a is one of the few Adventist scientists consequence of Satan, evil and who has addressed some of these is- the fall of humankind. Thus it sues. Essentially he presented some might be claimed that they are mixed models for discussion, explor- consistent with such scriptures as ing various aspects of pre-fall death. Rom 5:12-19 and Rom 8:22. Brand touched on the vexed issue of • They may be said to reflect cores in Eden, taking the view God’s omniscience. Perhaps that it “does not seem reasonable to an all-knowing God who could suggest that they accumulated and foresee man’s fall might be lasted forever.” He also noted that expected providently to build the fruits which comprised part of in mechanisms which could be the original diet begin as flowers and adapted for dealing with this become edible only after the flower contingency when it occurred. petals die. A similar picture, he pointed out, emerges with grains. Brand also • These views are entirely referred to the dung beetle, a species compatible with all the known whose whole life cycle appears to be functions and instances of cell designed around recycling. Although death, not just those related to Brand did not commit to any one the immune system, as in the position he suggested that a variety of Post-fall model. They are not views are compatible with scripture, embarrassed by apoptosis in including one which acknowledges embryological development, in the pre-fall existence of the death of cell death processes within the plant cells, the generation of some skin and gut, or by the death waste products and some recycling of of flowers occasioned by the

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production of Edenic fruit. associated with fiat creation. Evolu- tionary views have not found many They also suffer from some inadequa- adherents among literalist readers of cies. scripture. Within this context most • In admitting that programmed Seventh-day Adventists authors have cell death was part of the original clearly distanced themselves from such creation it could be argued positions33, although there have been that adherents of composite some exceptions.34 Certainly, some views have fundamentally evolutionary version is accepted by compromised on this question other conservative Christians and very and commenced travel on a widely so throughout at “slippery slope” which can only large.35 Accordingly, it is appropriate end in attribution of all death to also to critique this view in the context God Himself. of the creationist models with respect to the consistency of the manner in • Such views are also faced with which it contextualises and explains the serious challenge of deciding programmed cell death. at what level in the hierarchy of organisms death becomes Adherents of theistic would attributable to “sin” and hence regard such models as being successful morally “bad”. in explaining significant features of cell death. These successes are mainly • They also struggle to adequately in the scientific area. understand the function of the Tree of Life, for similar reasons • The presence of simple cell to those outlined under the Post- death mechanisms in quite fall model. primitive organisms, with a gradual diversification • Those committed to a high view and increase in complexity of the prophetic authority of apparent as one approaches Ellen White may understandably the higher taxonomic levels, argue that such views are in clear is consistent with prevailing opposition to the picture which evolutionary views suggesting emerges from her pen in so that mechanisms controlling cell many places. death appeared early in cellular 10.3 Evolutionary Models for the evolution, both within single- Origin of Programmed Cell Death celled (cells lacking Theistic evolutionary models suppose a membrane-bound nucleus) that God either instigated or allowed and during the development the gradual development of life over a of eukaryotic (cells with a much longer period than that usually membrane-bound nucleus)

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multicellular organisms. Once probability for the survival of the it occurred as a vital part of fittest as regards the population defence it became part of the as a corporate whole, which larger evolutionary process in could be expected to promote multicellular organisms, and rapid genetic evolution of the subsequently proliferated.36 species. Recycling of nutrients from apoptotic cells would also • The presence in eukaryotic maximise energy efficiency and cells of genetically independent thus contribute to the health of mitochondria may not represent the overall colony. the optimal design one might expect in terms of a creation • The continued development model. Evidence suggests that of colonial and multicellular the latter became intracellular organisms is favoured by symbionts very early in mechanisms of programmed eukaryotic evolution37. It is also cell death. This is so both for true that once this took place the protection via phagocytosis and significant interactions between in moulding the colony, in the mitochondria and the rest of same way as embryological the cellular structure, with their development is dependent on nuclear genetic dependencies, apoptosis. would have dramatically • The evolution of immunity both increased the possibilities for in its primitive innate form and adaptive mechanisms such as in more adaptive forms requires apoptosis and other forms of apoptosis to allow for clonal programmed cell death. selection, an instance of natural Additional successes may be suggested selection at the cellular level. by the arguable survival advantage This model presents significant prob- of organisms that have incorporated lems for many conservative Christians, mechanisms of programmed cell death. mostly at the theological level. • In the case of prokaryotic • The acceptance of any organisms mechanisms of evolutionary model for the origin programmed cell death in the of life implies a long history circumstance of limited nutrients for life and most likely for man would allow for rapid as well. Such a of the population of organisms is seen by many Christians as by maximising nutrients to those incompatible with the biblical most adapted to utilising them.38 account. Individual suicide of “stressed”' cells provides maximal • implies a long

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history of ecological systems to present new challenges to Christian involving differential survival understanding. The integration of and death or at least differential programmed cell death into this discus- fertility in the presence of sion provides an excellent example of mortality. Such a picture of this process. “nature red in tooth and claw”, QUESTIONS prior to any possibility of a 1. Do you see the programmed human fall and over such long death of cells as being “death” in periods, is again seen by many the same sense as that conveyed Christians as incompatible with by the death of a simple both the creation story and the organism, such as a worm? biblical picture of God. What about the death of a human • According to this model being? Are there different it is clear that immortality dimensions of death, or is there for humans can only result perhaps a , from a superposition on the lower levels of which may have natural order by God or a total regularly taken place in Eden reconstruction by Him of that without violating traditional order. Christian understanding of pre-fall perfection and 11. CONCLUSION deathlessness? It is clear that none of the models outlined are without problems. This 2. Which of the models discussed might be expected whenever finite at the end of this paper do you human attempt to probe events regard as representing the most deep “in the beginning”. Even greater honest compliance with firstly, explanatory difficulties might be an- the biblical data and secondly, ticipated in the event of there being an the scientific data? Do you evil power, loosed in before sense any tensions between your the world began, and functioning as a preferred biblical position on, “fifth-column” causal agent in God’s for example, the origin of death, creation. Just such an understanding and your preferred scientific does appear to permeate the Bible and position? How should a modern it is also present in all major Christian Christian best resolve such traditions since apostolic times. difficulties?

As new data emerge from scientific 3. Are you surprised to learn enquiry dialogue will continue be- that death mechanisms are tween scientists and interpreters of programmed into life forms at the Bible. This conversation, now the most fundamental level, i.e. hundreds of years old, can be expected the genome? Would you expect

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any programmed cell death at all the flaming sword which turned every in the New Earth? direction to guard the way to the tree of life” (Gen 3:2224,­ NASB). 4. Do you see the usefulness of the Bible as a modern guide to faith 3 “. . . and He will wipe away every as compromised in any way by tear from their eyes; and there will the fact that its writers obviously no longer be any death; there will no didn’t know much of what we longer be any , or crying, know about the natural world or pain; the first things have passed and the way life, for example, away'' (Rev 21:4, NASB)., “. . . On works? either side of the river was the tree of life, bearing twelve kinds of fruit, 5. Do you see any moral difference yielding its fruit every month; and the between the incidental death leaves of the tree were for the healing of a cell, as in necrosis, and of the nations. There will no longer programmed cell death? For be any curse; and the throne of God that matter, is there any and of the Lamb will be in it, and His moral difference between the bond­servants will serve Him” (Rev of a human 22:2, 3, NASB). being and an execution? 4 “For God so loved the world that REFERENCES: He gave His only begotten Son, that 1 “He will swallow up death for all whoever believes in Him shall not time, And the Lord GOD will wipe perish, but have eternal life” (John tears away from all faces, And He will 3:16, NASB). remove the reproach of His people from all the earth; For the LORD has 5 “And he ordered us to preach to the spoken” (Isa 25:8, NASB)., “The last people, and solemnly to testify that this enemy that will be abolished is death” is the One who has been appointed by (1 Cor 15:26, NASB). God as Judge of the living (‘quick’, KJV) and the dead” (Acts 10:42, 2 “Then the LORD God said, ‘Behold NASB). the man has become like one of Us, knowing good and evil; and now, he 6 “Only you shall not eat flesh with might stretch out his hand, and take its life, that is, its blood” (Gen 9:4, also from the tree of life, and eat and NASB)., “For as for the life of all live forever’− therefore the LORD flesh, its blood isidentified with its life. God sent him out from the garden of Therefore I said to the sons of Israel, Eden, to cultivate the ground from ‘You are not to eat the blood of any which he was taken. So he drove the flesh, for the life of all flesh is its blood; man out; and at the east of the garden whoever eats it shall be cut off’” (Lev of Eden He stationed the cherubim and 17:14, NASB)., “Only be sure not to

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eat the blood, for the blood is the life, sure and at temperatures above 100 and you shall not eat the life with the °C. Other examples are halophilic flesh” (Deut 12:23, NASB). organisms that live in the salt pans on salt lakes and only grow at NaCl 7 In Genesis 1:11 − 12, in which concentrations greater than 1M. Al- passage is depicted the creation of though such extremaphiles live at the vegetation, there is no mention of life. extremes of environmental conditions However, in vs 21, in connection with they nonetheless have a narrow range the creation of birds and fish, we find of temperature and environmental the expression “every living creature conditions that allow their survival that moves” (NASB). Furthermore, in and growth. Gen 9:3 we find God allowing “every moving thing that is alive” to be used 11 Kerr, J. F. R., Wyllie, A. H., and Cur- as food even as He had earlier given rie, A. R. (1972). Apoptosis: a basic humans the “green plant” as food. No- biological phenomenon with wide- where, so far as the authors could find, ranging implications in tissue kinetics. does the Bible explicitly speak of the Br J Cancer, 26, 239-257. “death” of plants, although there are 12 Yuan, J. and Kroemer, G. (2010). some confusing references, for exam- Alternative cell death mechanisms in ple, Eze 31:14. These points are often development and beyond. Genes & presented by writers within the Crea- Development, 24, 2592-2602. tion Science movement, such as John D. Morris – see http://www.icr.org/ 13 Clark, W. R. (1996). Sex and the article/1099/ (downloaded 6/6/2013). Origins of Death. New York: Oxford University Press, 31. 8 Timms, B. (2012). Seasonal study of aquatic in five sets 14 Kanold, P. O. (2009). Subplate of latitudinally separated gnammas in neurons: crucial regulators of cortical southern Western Australia. Journal development. Frontiers in Neuro- Royal Society of Western Australia, anatomy, 3,1-9. 95, 13-28. 15 Courchesne, E., Mouton, P. R., 9 See http://www.nobel.se/medicine/ Calhoun, M. E., Semendeferi, K., laureates/2002/presentation­speech. Ahrens-Barbeau, C., Hallet, M. J., html for a brief summary of the basis Barnes, C. C. and Pierce, K. (2011). of this prize. Neuron number and size in prefrontal cortex of children with autism. Journal 10 Extreme examples include the of the American Medical Association, thermophilic bacteria that live in deep 306(18), 2001-2010. sea vents where volcanic material is released into water. Here organisms 16 Medawar, P. (2013). Nobel Lecture: live at many atmospheres of pres- Immunological Tolerance. Nobelprize.

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org. Nobel Media AB. Web.3 Oct 25 Gen 2:7, 8. 2013. . ostrich; Job 38:39 & Ps 104:21− the 17 Doubling each time, an expansion young lions roar after their prey, and over 8-10 cycles will produce 28 (256) seek their meat from God; Job 38:41 & to 210 (1024) new cells for each initial Ps 147:9 − God feeds the ravens; Matt responder. 8:31, 32 − Christ sends the devils into the herd of swine which immediately 18 Clark, W. R. (1996). Sex and the drown; Luke 5:4-7 − the huge catch Origins of Death. New York: Oxford of fish. University Press, 93-95. 27 See Genesis 1-3; Isa 11:6-9; Isa 19 Lippens, S., Denecker, G., Ovaere, 65:25 and Rev 22:3. P., Vandenabeele, P. and Declercq, W. (2005). Death penalty for keratino- 28 White, E. G. (1908). Steps to Christ. cytes: apoptosis versus cornification. Washington, DC: Review and Herald, Cell Death and Differentiation, 12, 9. 1497-1508. Also Lippen, S., Hoste, 29 White, E. G. (1958). Patriarchs and E., Vandenabeele, P., Agostinis, P. and Prophets. California: Pacific Press, Declercq, W. (2009). Cell death in the 62. Possibly also reflecting a view skin. Apoptosis, 14, 549-569. of the of flowers in Eden 20 See for details Doctor, K., Reed, J., Ellen White reported an early vision Godzik, A., and Bourne, P. (2003). The in which she seemed to be in the New apoptosis database. Cell Death Differ, Earth. While plucking some flowers 10(6), 621-633. she cried out, “They will never fade.” (White, E.G. [First published 1882] 21 Levine, M. and Tjian, R. (2003). (1945). Early Writings. Washington, Transcription regulation and animal DC: Review and Herald, p18). A diversity. Nature, 424(6945), 147-151. similar statement by the same author 22 Fisher, H.J. (1987). The enigma of appears elsewhere. “This earth,... pu- ecology. Record, 92(11), 4-5. rified with fire, then ... will be much more beautiful. The grass will be 23 Provonsha, J. (2000). ‘Creation/ living green, and will never wither. Evolution Debate in Light of the Great There will be roses and lilies, and all Controversy’, in Hayward, J. L. (ed.), kinds of flowers there. They will never “Creation Reconsidered: Scientific, blight or fade, or lose their beauty Biblical and Theological Perspec- and fragrance'' (White, E.G. (1976). tives”. California: Assoc. of Adventist Maranatha. Washington DC: Review Forums, 303-311. & Herald, p355). 24 Gen 1:28.

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30 White, E. G, (1958). Patriarchs and reflect awareness of long-age and evo- Prophets. California: Pacific Press, 48. lutionary concepts explored in several of the essays in Hayward, J. L. (2000). 31 Rev 22:2. Creation Reconsidered: Scientific, Bib- 32 See Brand, L. (2003). ‘What are the lical, and Theological Perspectives. Limits of Death in Paradise?’, paper California: Association of Adventist presented at the South Pacific Divi- Forums., particularly those by Taylor, sion Faith and Science Conference at E. and Ritland, R.M. Also, although Avondale College, July 11-14. not explicitly espousing theistic evo- lution, the book: Bull, B., Guy, F. & 33 See for example: Ball, B. (ed), Taylor, E. (eds.), (2006). Understand- (2012). In the Beginning. Idaho: Pa- ing Genesis: Contemporary Adventist cific Press., Gibson, L. J., and Rasi, H. Perspectives. California: Adventist (eds), (2011). Understanding Creation. Today, sets up a framework which Idaho: Pacific Press., Clausen, B. allows theistic evolutionary options. and Wheeler, G. (2006). The Book of Beginnings. Hagerstown, MD: Review 35 See for example: Collins, F. (2006). and Herald., Roth, A. (1998). Origins: The Language of God. NY: Free Linking Science and Scripture. Hagers- Press., Colling, R. G. (2004). Random town, MD: Review and Herald. and Designer. IL: Browning Press., Falk, Brand, L. (1997). Faith, Reason and D. R., (2004). Coming to Peace with Earth History. Berrien Springs, MI: Science. IL: Intervarsity. and Peters, Andrews University Press. T. & Martinez, H. (2003). Evolution from Creation to New Creation: Con- 34 In his lecture, “Adventist Interpreta- flict, Conversation, and Convergence. tions of Genesis 1:1,2”, presented at Nashville, TN: Abington. Avondale College on May 14, 2011, Gerhard Pfandl acknowledged the 36 See Taylor-Brown, E. & Hurd, presence of this viewpoint with the H.2013). The first : a legacy statement, “The third view in our inherited by parasitic protozoans Church (Adventism) is theistic evolu- from ancestors. Parasites tion. As I indicated this morning for a and Vectors, 6,108. Also Ratcliff, W. number of years now C., Denison, R. F., Borrello, M. and has been held by several scientists and Travisano, M, (2012). Experimental theologians”. Pfandl mentioned Fritz evolution of multicellularity. PNAS, Guy by name. Later in his lecture this 109, 1595-1600. point was repeated: “Now, in addi- 37 Gray, M. W., Burger, G. and Lang, tion to the two views indicated in this B. F. (1999). Mitochondrial evolution. quote we find now also a third view, Science, 283, 1476-1481. theistic evolution, being promoted in the Church today.” These comments 38 See Koonin, E. and Aravind, L.

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(2002). Origin and evolution of eukaryotic apoptosis: the bacterial connection. Cell Death Differ, 9(4), 394-404., and Bridgham, J., Wilder, J., Hollocher, H., and Johnson, A. (2003). All in the family: evolutionary and functional relationships among death receptors. Cell Death Differ, 10(1), 19-25.

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