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Hi All

Arking maybe should be requested to review Walford's new CR book, based

on

the below. It is from a not yet in Medline review. CR forms much of the

discussion

of the review.

BIOLOGY, FRUIT FLIES, AND HUMANS: CAN EXTENDED LONGEVITY STRETCH FROM ONE TO THE

OTHER?

Review Essay by Arking

Gerontologist. 2005 Jun;45(2): 418-425.

The Biology of Death: Origins of Mortality, by André Klarsfeld and Frédéric

Revah

(translated from French by Lydia Brady). Cornell University Press, Ithaca, NY,

2004,

211 pp., $29.95 (cloth).

" How should we live our lives? " is an age-old question. " Should we actively seek

to

live a longer life? " is a more recent question now in the process of being

answered.

" How should we live an extended life? " is a newborn question just beginning to

be

addressed. Just as there is a multiplicity of views regarding the proper answer

to

the first question, so there are differing views regarding the latter two

questions.

While these divergent and competing views may be judged by some as an

inappropriate

cacophony, others may view them as an ongoing discussion and commentary on the

most

personal of all things—our lives and their shape. The three books reviewed in

this

essay ostensibly address the second question, although their authors' views can

be

interpreted as pointing towards some still-forming response to the third

question.

A Dinner Conversation: Death and Longevity

A stimulating dinner conversation is stated to have been the inspiration for The

Biology of Death: Origins of Mortality, by André Klarsfeld and Frédéric Revah.

They

have written a short but pointed summary of recent scientific findings regarding

death and longevity for the intelligent lay person. People tend not to dwell too

much on death except to believe that it must be a biological necessity, if only

to

justify the unfairness of that personal extinction that awaits us all. This book

is

the authors' response to that common but flawed perception.

Originally published in French in 2000, this translation captures the flavor of

the

French writings on this topic. I for one thank Lydia Brady for her efforts,

which

make only too clear the loss of access to all those untranslated writings which

afflict us unsuspecting monolinguists. The authors start off with a concise

overview

of prior biological concepts of aging and death, winding up with August Weismann

and

Elie Metchnikoff and the beginnings of the evolutionary concepts of aging.

Following

a discussion of the demographic methods of measuring longevity (complete with

charts

and tables), they engage in a comparative examination of the rapid and gradual

modes

of aging. Only then do they segue into a continuation and elaboration of the

evolutionary theory of aging, one in which they weave the essence of the

theoretical

arguments and their supporting empirical data into an engaging story (with

minimal

jargon) that culminates in the disposable soma theory and the purposelessness of

death.

In a mere 30 pages Klarsfeld and Revah summarize the current state of our

knowledge

of the mechanisms underlying aging in a manner such that a careful lay reader

could

hold her own at that dinner table conversation. Following a discussion of

apoptosis,

they delve into an examination of the ways in which one might significantly slow

the

aging process. In the few years since Klarsfeld and Revah wrote this

well-crafted

slim volume in French, our knowledge of aging mechanisms and potential

interventions

has qualitatively increased. So this portion of the translated text is the most

dated (although the French version must have been quite current when published).

They recap the conversation by concluding that " the priority of living organisms

cannot be to devote all of their efforts to their own survival; they must also

keep

resources to reproduce, to transmit their genes. Aging and natural death arise

from

this compromise, not automatically by a direct link but through the work of

natural

selection; they are simply particularly adverse side effects, from a human point

of

view " (pp. 193–194). In this, they agree with the now-accepted view based on the

prior conclusions of scholars such as Rose (1991), Kirkwood (1987),

(1957),

Medawar (1952), and Bidder (1925, 1932) regarding the role of reproduction in

bringing about the lack of somatic maintenance, which is the ultimate cause of

death.

The evolutionary approach to understanding aging is the only concept that has

allowed us to construct a coherent and empirically proven explanation as to why

organisms possessed of an elaborate synthesis and repair apparatus should

inevitably—and otherwise inexplicably—fail to maintain themselves and so

inexorably

lose function, decay, and die. Klarsfeld and Revah's dinner companions must have

been fascinated by their conversation, as much by the light it shed on the

problem

of death as by the method of its analysis. As Rose has well written

elsewhere, " And for those who would know and understand the long story of life

on

earth, Darwinism is the great searchlight in the darkness. For the modern world

to

go on without Darwin's Spectre would be to lose our way in a twilight of the

mind "

(Rose, 1998, p. 211).

Not all celebrate the intellectual triumph of this materialist approach to the

deep

questions of life, for many are uncomfortable with the absence of a

compassionate

creator. The supposed consolations of death and finitude have been put forth in

moral terms, most notably by Leon Kass (2004). (For a specific rebuttal to his

arguments see Arking, in press.) They have also been expressed in supposedly

biological terms, the latter exemplified by Klarsfeld and Revah's summary of the

precepts of the Thanatology Society, which posits that " ... the death of

individuals

ensures not only the perennity of the species but also its rejuvenation: it is

hence

not only a necessity but a benefit " (M. Marois, p. 192). There is some truth to

this

statement, involving the calculation of Darwinian fitness in changing

environments.

But we now realize that an increase (or a decrease) in the frequency of some

allele

which we pass on to our offspring has no moral meaning. It offers us no

beneficial

recompense for our death. " What compensation would the immortality of the

species

bring to the individual stalked by death? ... What good does it do me that the

future world will survive if I won't be there? " (V. Jankelevitch, p. 193).

This individualist and materialist rejection of mortality's alleged benefits was

not

only voiced by others, but is becoming more widespread in our society as

evidenced

by a recent article by Kaufman, Shim, and Russ (2004). They argue that

biomedical

improvements make the improbable treatment routine, change the expectations for

the

quality of life at older ages, and bind hope to the normalization of

life-extending

techniques. In effect, the alleged benefits of death are being refuted by every

individual who takes the new drug or undergoes the new surgical interventions

that

modern data-based medicine suggests will likely extend his or her life.

It seems as if ordinary people are exerting their autonomy by opting for life

rather

than for death. The intellectual validity of their optimistic actions is

supported

by the prolongevity philosophy put forth by Overall (2003, 2004). A

materialistic explanation of human origins in an uncaring universe will likely

resonate only with that subset of people who can be comfortable with the cold

comfort of logic and who can derive " ought " from " is " (see Bronowski, 1956). In

contrast, those desirous of a purposeful universe overseen by a compassionate

creator may consider such a view as devoid of meaning and incapable of giving a

moral coherence to life. The political future of prolongevity interventions will

likely rest on our ability to truthfully present to both groups the information

necessary for them to individually view such interventions as supportive of life

and, thus, of their own philosophy.

Prolonged Senescence

In the current societal state of affairs, life expectancy is being extended by

adding years onto the latter part of the life span. The social and ethical

discussions swirl about this one strategy only, although other and better

strategies

are available (as will be discussed below). This current strategy works only

because

social organisms may receive enough support from their group to enable them to

survive past the time when they would otherwise have been able to survive on

their

own. Such supportive behavior was selected for in humans by both cultural and

biological means as Crews (see below) points out in much detail. But

such

amelioration, even though it is an intrinsic part of our species' life history

strategy, does not defer senescence but lengthens it. What might we expect to

happen

if we continue to follow this strategy of extending senescence? There is good

news,

and there is bad news (Baltes & , 2003). The good news is that people in

the

65–85 age bracket (which constitute about 12.8% of the population of developed

countries) are much healthier and more productive today than they were in the

past,

and they have high levels of physical and mental well-being. Moreover, there is

still substantial latent potential for better mental and physical fitness among

them. For these reasons, people in the 65–75 age bracket have been termed the

" young–old " in recognition of their ability to maintain function into what was

traditionally considered the frail and morbid years of being " old. " Many of

these

young–old adults could continue to contribute to society and enrich it by

continuing

to work, by developing new social roles, and by contributing in various ways to

the

resources needed for their own medical care. In fact increasing numbers already

do

so, and our society will likely change for the better as this trend continues.

The

bad news is that the " oldest–old, " the people in the 85–100 and older age

bracket

(which constitute about 1.5% of the population in developed countries), are not

in

as good a condition. Many have some sort of functional physical or cognitive

loss

and have also lost their social contacts as spouses and relatives and friends

die.

These oldest–old adults are, for the most part, lonely women, most of whom will

die

alone in a hospital or nursing home. This group may not be numerically large

now,

but their numbers will substantially increase if we just maintain our current

practices of biomedical intervention at the end of life. They are a harbinger of

our

future.

Such a prospect should give even an optimist pause, for the joys of living long

are

nullified by the loss of dignity and control suffered by the oldest–old. If all

that

our vaunted knowledge of the biology of aging can do is to increase the odds

that

one would live into the oldest–old group without devising any way to buffer one

against the inevitable functional losses seen in this group, then perhaps it

would

be better for biogerontologists not to continue their research. It is enough to

make

one wonder whether there should be limits to life. Kaufman and colleagues (2004)

point out that " A price is paid for hope and expectation invested in biomedical

technique " (p. 737), and that price seems to be one of deficits, disappointment,

and

despair when the hopes of " eternal " health do not materialize, as indeed they

cannot.

Delaying the Onset of Aging

But there is another strategy available, and it grows out of modern

biogerontological research. One of the major purposes of using laboratory models

for

research is to discover mechanisms and responses that may not be obvious in

humans

but that can be brought to light by animal studies. Should a new or unusual

response

be found in mice or primates, then we should consider the possibility of it

occurring in humans and the implications. In Methuselah Flies: A Case Study in

the

Evolution of Aging, by R. Rose, Hardip B. Passananti, and Margarida

Matos,

the very first research paper is a reprint of an earlier report by Rose on the

evolution of postponed senescence in Drosophila (Rose, 1984). This paper was

published simultaneously with the independent report of Luckinbill, Arking,

Clare,

Cirocco, and Buck (1984) showing the same effect. Much work has been done since

then

elucidating the mechanisms underlying this form of extended longevity, as the

new

book by Rose and colleagues amply demonstrates.

The work that my colleagues and I have since done on Drosophila longevity bears

on

the development of this alternate strategy. We reported that aging in the Ra

strain

of wild type flies is rather complex, being characterized by at least three

different extended longevity phenotypes, each of which was induced by specific

stimuli and had different demographic mortality and survival profiles (Arking,

Buck,

Novoseltsev, Hwangbo, & Lane, 2002).

The first longevity phenotype (Type I) is a delayed onset of senescence which

leads

to a significant increase in both the mean and maximum life span of the

experimental

strain. Normally, mortality rates increase as a population ages. It turns out

that

an extension in life span may be brought about either by an age-independent

reduction in the initial mortality rate observed in the youngest members of a

population or by an age-dependent reduction in the rate of increase in the

mortality

rate with age. The former case results in the same rate of increase in the

mortality

rate as a function of age, but each age cohort is much healthier than is its

comparable age cohort in the relevant control population because their starting

mortality was so much lower. The net effect is an increase in longevity due to

increased health. The latter case results in the population having the same

initial

mortality rate as the control, but this is coupled with a lower rate of increase

in

the mortality rate with age. As a result, each age cohort is aging slower than

is

its comparable age cohort in the control population. The net effect is an

increase

in longevity due to a slower rate of aging. Demographic analyses of our

long-lived

La strains shows that they are using the second method, as their rate of aging

is

reduced by 33% relative to normal-lived controls.

This is not true of the other two longevity phenotypes observed in these Ra

strain

flies. The second (Type II) is characterized by an increased early survival

which

leads to a significant increase in mean but not in maximum life span. The third

(Type III) longevity phenotype is an increased later survival which leads to a

change in the maximum but not in the mean life spans. There is no significant

alteration in the rate of aging in either of these alternative longevity

phenotypes,

although there is a transient decrease in mortality in the young (Type II) or

the

old (Type III) individuals.

Moreover, empirical evidence demonstrates that not only do multiple longevity

phenotypes exist but that each of them may be induced by a variety of stimuli,

some

of which may interact in an additive manner. The Type I delayed onset of

senescence

phenotype may be induced in flies by (a) caloric restriction (Pletcher,

Mac,

Marguerie, Certa, Stearns, Goldstein, et al., 2002), (B) the down regulation of

the

insulin-like signaling pathway (Clancy, Gems, Harshman, Oldham, Stocker, Hafen,

et

al., 2001; Tatar, Kopelman, Epstein, Tu, Yin, & Garofalo, 2001), © up

regulation

of the antioxidant defense system (ADS) plus altered mitochondrial properties

(Arking et al., 2002), (d) different alterations through pharmaceuticals of the

patterns of adult gene expression (Kang, Benzer, & Min, 2002; Zhao, Sun, Lu, Li,

Chen, Tao, et al., 2005), and (e) alteration of steroid hormone levels (Simon,

Shih,

Mack, & Benzer, 2003). These different stimuli in flies probably exert varied

specific effects, but all seem to lead to high levels of somatic maintenance and

thus to a significant delay in the onset of observable loss of function. The

mechanisms likely involved are discussed elsewhere (Arking, in press; see also

Rose

and colleagues, as discussed below).

Furthermore, this diversity of longevity phenotypes is not restricted to

insects.

All three phenotypes are known in mice. However, only the three longevity

phenotypes

of Types II and III are known to occur in humans. The mouse data, when combined

with

the conservation of aging mechanisms across species (see below), strongly imply

their parallel existence in humans. There is every reason to believe that the

Type I

phenotype can be expressed in primates as a delayed onset of senescence. For

example, it certainly appears as if the Type I phenotype is expressed in

calorically

restricted macaque monkeys (Roth, Lane, Ingram, Mattison, Elahi, Tobin, et al.,

2002) and humans (Fontana, Meyer, Klein, & Holloszy, 2004; Walford, Mock,

Verdery, &

MacCullum, 2002). An underappreciated outcome of the past century of biological

research is the discovery that fundamental cellular and homeostatic traits are

common to almost all species and are very often controlled by the same basic

molecular mechanisms. Evolution may be said to be frugal in its invention of new

molecular mechanisms but extravagant in the number of species employing them.

Against that conceptual background, these findings almost certainly imply the

existence of an evolutionarily conserved longevity mechanism and its

corresponding

longevity phenotype. If so, then it should be possible for us to induce the

delayed

onset of senescence phenotype in humans using stimuli other than a 30% reduction

in

caloric intake. Such efforts are underway in various labs and biotech companies

(Hadley, Lakatta, on-Bogorad, Warner, & Hodes, 2005).

Historically, clinical therapeutic efforts focused on alleviating the

symptomatic

effects of underlying age-related diseases. The increase in our knowledge has

fostered recent clinical efforts to more effectively treat the underlying

chronic

conditions. The effect of these efforts has been to progressively decrease the

mortality of senescent adults over the past few decades. That strategy keeps us

alive longer than was the case in the past, and I am personally grateful for it;

but

it does not extend the healthy span of our lives. The failure of humans so far

to

express a Type I delayed onset of senescence phenotype may be the result of not

applying an appropriate stimulus to our bodies rather than being the result of

an

intrinsic inability of our bodies to respond. With the appropriate stimulus we

would

be asking our bodies to evoke a response which is already built into our genome.

This is by definition a natural response. Such a natural evocation of an innate

response would not alter any essential aspect of our human nature.

Pharmaceutical

interventions have been proved in principle on the basis of work reported with

worms

and flies and mice. The current goal of the advanced research into the biology

of

aging is to identify and characterize pharmaceutical inducers of the Type I

delayed

onset phenotype in mammals, eventually including humans. The success of these

efforts will likely weaken the current ethical arguments against extended

longevity,

based as they presently are on decreasing the mortality of senescent adults. We

will

pick up this point after we examine the book by Crews below.

As noted above, Rose's 1984 seminal paper helped to open the modern phase of

biogerontological research. What of the book in which it is recapitulated?

Methuselah Flies: A Case Study in the Evolution of Aging is a most unusual

volume.

It is not the usual festschrift, lovingly assembled at the end of a mentor's

career.

Nor is it the usual edited assemblage of varied individual contributions

hopefully

focused on some common theme. This is instead a deliberately crafted compilation

of

29 scientific papers by Rose and his colleagues, all of which are

focused on

the analysis of his well-known normal-lived (B) and long-lived (O) strains of

Drosophila melanogaster. Note that Rose has authored a minimum of 40

peer-reviewed

articles on these strains over the past 20 years, in addition to a large number

of

review papers, which makes it apparent that this is not a mere pastiche of his

reprint files. This book represents a deliberate overview of the knowledge

gained

from the past 20 years of work with these long-lived fruit flies. This

organization

of their knowledge resulted in six lacunae, and so six papers were written

especially for this volume. They underwent an independent peer-review process

under

the guidance of Margarida Matos, which resulted in the rejection of one of them.

The

remaining 5 newly vetted articles and the 24 previously published articles are

organized around six general themes (each preceded by a short introductory

section)

as follows:

Creation and long-term evolution of Methuselah flies;

Stress, resistance, physiology, and aging;

Reproduction, nutrition, and aging;

Genetics and molecular biology of Methuselah flies;

Reverse evolution of Methuselah flies;

Aging, development, and crowding.

This is not a book for the uninitiated lay person, but it is one with which an

experimental gerontologist can curl up. I found the introductory essays useful

and

the compiled papers convenient to compare. The goal of the book is to emphasize

the

value of the research on the array of long-lived flies created by Rose and his

collaborators. The perception of that value has been eclipsed in recent years as

work done with single-gene mutants of yeast, nematodes, flies, and mice has

allowed

the identification and characterization of the genetic and molecular pathways

regulating the organism's stress resistance and, hence, longevity. This book is

an

effort to bring attention back to the focused work done by Rose and his

collaborators through a convenient collection. Much of that value stems from the

fact that few labs have focused such a variety of experimental techniques on one

set

of related long-lived and control animals.

So what has this work taught us about the mechanisms underlying this delayed

onset

of senescence phenotype? Other than the creation of the stocks of fruit flies,

which

was instructive and much imitated, the work done on topics 2, 3, and 5 (listed

above) yielded the most useful set of data and concepts. It had long been

thought

that long-lived animals reworked their physiology so as to increase their

caloric

investment in somatic maintenance at a cost of early fecundity, only to mobilize

their resources for reproduction in later life. The papers in these three topics

adduced much of the data that supported that early assumption. Long-lived

animals

really are physiologically different animals, and we should not be surprised

that

their reengineered energy metabolism allows them to age slower. The detailed

studies

on nutrition and stress resistance helped to define the major parameters

defining

these two important variables which directly affect the onset of aging and

senescence. But no one approach is perfect, and the work done with selected

strains

of fruit flies summarized in part 4 of the Rose volume gives no hint of the

powerful

effects on longevity of particular single-gene mutants, especially those

involved in

the insulin-like signaling pathway and its control of stress resistance (Helfand

&

Rogina, 2003; Kenyon, 2005). The population studies of topic 1 listed above

suggested a large number of genes were involved in aging, which is true; but the

inability to discern the major reworking of the animal's physiology brought

about by

the altered expression of individual key regulatory genes and particular

pathways

led to the underappreciation of the single-gene approach.

However, it must be pointed out that the success of the molecular genetic

approach

depended on the prior existence of Riddle, Swanson, and Albert's (1981) classic

analysis of certain stress resistance pathways in the nematode. These were given

significance for aging by the later serendipitous finding that certain of these

mutants defining a certain type of stress resistance were long-lived (Kenyon,

Chang,

Gensch, Rudner, & Tabtiang, 1993; Larsen, Albert, & Riddle, 1995) and that one

of

the key mutant genes carried the information required to synthesize a key

component

of the insulin-like signaling system. The upshot was the connection of a

regulatory

system extraordinarily well described in mammals (the insulin-like signaling

system)

but not in worms or flies, with the well-developed concepts of extended

longevity in

these invertebrates. The connection of two such different phenomena made no

sense

unless one understood it to imply the existence of an evolutionarily conserved

set

of regulatory mechanisms modulating aging in invertebrates and metabolic

regulation

in vertebrates. The hypothesis that aging and metabolic regulation were

connected in

both species was rapidly tested and proven correct, and the concept of

manipulating

aging came to the minds of some (e.g., Guarante, 2003). Fortuitous findings from

independent laboratories seem to have trumped a more planned approach, at least

in

this sphere.

Evolutionary Biology and Cultural Factors

But how much of this is relevant to us? Humans are not flies, worms, or just

featherless bipeds. We share about 98% of our genes with the chimpanzee, who is

our

closest relative, but we are not just bigger chimps. Crews makes this

very

clear in Human Senescence: Evolutionary and Biocultural Perspectives, his

excellent

examination of " the evolutionary biology of human senescence from an

evolutionary

and biocultural perspective " (p. vii). One cannot reliably translate the

laboratory

findings on laboratory model organisms to humans without a full understanding of

the

nature of our evolutionary differences compared to other hominids and primates

or of

the huge but hitherto underappreciated role which our cultural adaptations

(e.g.,

intergenerational transfers of resources) play in allowing us to be described as

being very long-lived hominids with a generally high (albeit variable) level of

postreproductive vitality not seen in other forms. Crews' slender but

informationally dense book does an excellent job of linking the findings of

human

anthropology to those of biogerontology (and vice versa). Perhaps this book can

be

viewed as a continuation of the Klarsfeld–Revah dinner conversation, only now

focusing on the nitty-gritty details that differentiate fact from fiction.

Following

two introductory chapters, Crews moves into detailed discussions of human

variation

in growth and life history (chapter 3), as well as chronic diseases, risk

factors

and senescence (chapter 4), and thence segues into a discussion of human life

span

and life extension (chapter 5), before ending with an overview and perspective

of

all the material (chapter 6).

Although Crews points out that we are unarguably different from other primates,

there are arguably no unique human qualities. Instead our uniqueness stems from

an

unusual array of interrelated adaptations that improve the adaptability of each

quality. These include bipedality, large brains, high visual dependence, verbal

communication, culture, manual dexterity, complete infant dependency, growth and

development patterns quite different from those of our primate cousins, and long

life. Consider our growth patterns. Our intrauterine growth period of 9 months

is

comparable to that of chimpanzees and gorillas, but it results in big-brained,

comparatively large—but helpless—infants who cannot even cling to their mothers

but

do have " a well-developed ability to be heard, after which they are continually

held

to the breast by the free upper limbs of their mothers " (Crews, p. 85). Even

though

our infants are twofold larger at birth than are our primate cousins, they are

weaned at half the age. The human postnatal growth period is unlike those of

other

primates, which have neither midchildhood nor adolescent growth spurts nor even

such

life stages. Compared to chimpanzees, we have a much slower physical development

both in utero and postnatally, coupled with a comparatively slow pattern of

neural

development. For example, our brains attain about 25% of the adult weight by

birth,

attain adult weight by around 10 years, but continue developing until we are 20

years old, at least. We become sexually mature at about age 14, much later than

do

great apes (around 7–10 years), but significantly before we are fully developed

(around age 20), and much parental distress follows from that asymmetry. All of

this

developmental discussion is especially interesting because of the comparative

format

in which Crews has placed it.

As Crews points out, longevity in modern humans did not evolve directly but

rather

is secondary to the evolutionary trends listed above. Culture, communication,

and

language allow us to maintain relatively constant microenvironments and thus

play

crucial roles in our reproductive success. These biocultural responses

restructure

our genetic variation and alter existing gene–environment relationships,

allowing

long-lived, postreproductive survivors to contribute to the success of their

children and extended kin, thus further enhancing the longevity-favoring aspects

of

these biocultural adaptations. It is this biocultural adaptability that makes

human

life history different from other mammals. This phenomenon makes some question

whether the extended longevity discussed above (e.g., the Type I delayed onset

of

senescence) can be reasonably translated from laboratory data to the human

condition, or whether this scenario is just another snake-oil salesman's fairy

tale

of forbidden hope. In his discussion of this question, Crews rightly emphasizes

the

complex differences between humans and other hominids, particularly the fact

that we

are already a very long-lived species with an unexpectedly long postreproductive

life, often significantly longer than our reproductive life. But because our

history

has already made us long lived, then perhaps we are already at the limits of our

innate longevity; and if so, then perhaps the longevity interventions generally

operative in simpler laboratory model organisms will not be operative in us. The

tricks that let laboratory animals live long may not be effective in us if they

have

been superseded by our unique biocultural adaptations. There is no conclusive

answer

yet available to this question. So Crews leaves the issue formally open, as

indeed

the data would dictate that he should. But after reviewing the evidence, he does

conclude that " According to all available evidence, calorie restriction

instituted

during childhood should retard [senescence] and increase average life span for

humans .... [it] may represent an untapped source for human life extension ....

[N]utritional restriction appears to be humankind's best hope for a fountain of

youth " (pp. 209–210). I agree with him and wish to add some recent evidence to

his

argument.

The basic mechanisms underlying the senescent mechanisms that eventually do us

in

are not unique to us but rather common to all cells. Senescence is the result of

a

cell-based response to the body's nutritional status (de Cabo, Surer-Galban,

Anson,

Gilman, Gorospe, & Lane, 2003). Cell-level mechanisms and pathways are highly

related across species and often exhibit high levels of functional and

structural

homology. Interventions acting at the level of the individual cell which delay

the

onset of these senescent mechanisms should be effective across species because

they

are not directly affected by the peculiar higher level biocultural adaptations

which

Crews has described in some detail. The loss of function which underlies

senescence

mostly (but not entirely) involves the direct and indirect actions of oxidative

damage—a sort of biological " rusting " of the subcellular structures. It is often

accompanied by a slow but deleterious accumulation of damaged proteins—a sort of

" junk accumulation. " This rusting and junk accumulation process is opposed in

the

model laboratory organisms by a conserved assortment of stress resistance

mechanisms

and molecules. These resistance mechanisms can be induced by a number of

molecules

acting more or less directly on the cell (Marsh & , 2004; Morley &

Morimoto,

2004) in a cooperative manner ( & Lithgow, 2003). The experimental facts

presented in those reports form the basis for our present understanding of the

events that cause our cells to transit from a state of high function to a state

of

lower and decreasing function (see Arking, in press). But can we extrapolate the

data? Do our cells react to stress in the same way that other animals' cells do?

Although not yet complete, caloric restriction studies on macaque monkeys

clearly

show that this intervention has altered their physiology such that they appear

to be

losing function significantly slower than their normally fed cohorts (Roth et

al.,

2002). The National Institute on Aging (NIA) has recently initiated a study of

the

effects of caloric restriction on humans, but those data will not be known for

some

years yet (Heilbronn & Ravussin, 2003). However, some individuals have

voluntarily

put themselves on a calorically restricted diet, and a recent review of their

medical records suggests that humans likely undergo the same physiological

changes

seen in calorically restricted monkeys (Fontana et al., 2004). Among individuals

in

good health but not undergoing any known anti-aging intervention, the physiology

of

those who lived longer than the median age closely resembles the physiology of

the

calorically restricted macaque monkeys (Roth et al., 2002). Female athletes

undergoing intensive physical exercise often cease menstruating, an indication

that

we still retain the cellular mechanisms whereby the body can shift from a

reproductive mode to a somatic maintenance mode under stress conditions not

suitable

for reproduction. Given these findings, it is not an unreasonable speculation to

believe that longevity interventions that enhance our cells' abilities to resist

various stresses, and thus to delay the onset of senescence, stand a reasonable

chance of forming the basis of a pharmaceutical prolongevity intervention that

will

be effective in humans.

It is likely that such interventions will one day induce the appearance in

humans of

the " missing " Type I (delayed onset of senescence) phenotype. The social

consequences of this intervention are obviously speculative but might well be

rather

different from what might be expected at first flush. Because such an

intervention

would be adding healthy years to presenescent individuals, it is open to none of

the

philosophical criticisms directed towards the biomedical extension of senescence

(Kass, 2004).

These three books were interesting and informative to me, and I recommend them

to

all gerontologists and interested observers. The choice of which text you choose

might well depend on your familiarity with modern biology and/or your interest

in

humans as opposed to fruit flies. But each of these texts contains an

interesting

tale told by masters of their craft, and they are worth the reading.

Those individuals who lived throughout most of the 20th century often marveled

at

the changes they witnessed and the advances in the physical sciences that led to

the

inventions that transformed their world. There is a reasonable chance that the

advances in the biological sciences that will take place in this 21st century

will

equally transform the world, and do so in ways that perhaps seem unlikely to us

today. Now there is a topic for an interesting dinner conversation—and perhaps

another book.

Al Pater, PhD; email: old542000@...

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