Ashdin Publishing
Journal of Evolutionary Medicine
Vol. 5 (2017), Article ID 235975, 11 pages
doi:10.4303/jem/235975
ASHDIN
publishing
Research Article
THEORY
The Evolving Opportunistic Pathogen Communities on Host Individuals
and the Evolution of Host Aging
Ulfat Baig, Akanksha Ojha, and Milind Watve
Indian Institute of Science Education and Research, Dr. Homi Bhabha Road, Pune, Maharashtra 411008, India
Address correspondence to Milind Watve, milind@iiserpune.ac.in
Received 24 February 2016; Revised 4 April 2017; Accepted 6 April 2017
Copyright © 2017 Ulfat Baig et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Abstract In the coevolution of host and the associated oppor-
tunistically pathogenic microbiota, the microbiota has an advantage
of a smaller generation time and thereby faster evolution. Sexual
reproduction by the host is hypothesized to be the hosts’ evolutionary
counter-strategy. We propose further that the ticking clock of the
evolving microbiota influences the evolution of host aging. Modeling
these dynamics shows that if transmission of microbes has a small
to moderate vertical or kin-biased component, early aging can
evolve in the host. Host genotypes with shorter longevity are more
likely to escape pathogen evolution thereby getting a selective
advantage for their progeny when risk of infection is high. As parasite
communities are ecologically and evolutionarily dynamic, hosts can in
response evolve plasticity in aging. The model shows that a genotype
which activates aging or death pathways in response to threshold
parasite colonization gets a selective advantage whenever there is a
nonzero kin transmission bias. The hypothesis is compatible with
classical hypotheses for aging. We make many predictions testable by
epidemiological, comparative or experimental methods.
Keywords parasites transmission; evolution of lifespan; kin selection;
lifespan; life history
1. Introduction
Whether aging is inevitable or is a trait specifically evolved
under some kind of selection is an old debate. One school of
thought assumes aging to be inevitable and beyond the reach
of natural selection [1]. Many of the processes implicated
in aging such as accumulation of somatic mutations,
protein aggregation, oxidative damages or telomere
shortening might be biochemically and thermodynamically
inevitable [2]. However, since mechanisms of repair,
replacement, throwing away or asymmetric segregation
of damage exist, inevitability of mechanisms does not
necessarily mean inevitability of the process. Germ line
cells in higher organisms appear to perpetuate without any
signs of aging [3,4,5,6,7]. Since every new generation
begins with a single celled zygote, mutations in germ line
cells are directly subject to natural selection, which prevents
accumulation of deleterious mutations; therefore germ line
cells do not seem to age. Organisms such as hydra and
many plants that reproduce vegetatively do not show any
obvious aging [4,5,8,9,10,11]. In some species, viability
or reproductive capacity actually increases monotonically
with age [11, 12]. If decline in viability is not universal with
aging, it is worth asking why and how it declines in some
organisms including humans.
For a long time, bacteria that divide by binary fission
were believed to be immune to aging. The demonstration
of signs of aging in bacteria even during exponential
growth [13] changed the perspective. However, asymmetric
damage segregation, which is responsible for bacterial
aging, has been shown to be both plastic and evolvable [14].
Bacterial aging provides many other grounds to doubt the
inevitability of aging. Although some cells in a growing
clone of bacteria undergo senescence, the clone as a whole
grows with continued rejuvenation. A process that works
for clones of bacteria should be possible, in principle, for
multicellular organisms which are essentially clones of the
zygote.
Is aging inevitable after a germ-soma division of cell
lines? Even if one assumes that aging is inevitable for
a somatic cell, it does not necessarily imply that it is
inevitable for an organism. A multicellular organism may
program the cell turnover so as to replace the old cells in
a balanced manner and thus escape organismal senescence
indefinitely if resource provisioning is adequate. On the
other hand, senescence of a system is theoretically possible
even if its components are individually nonaging but
irreplaceable [15]. Therefore, cellular senescence is neither
necessary nor sufficient to explain organismal senescence.
Since inevitability of aging is theoretically inadequate
and empirically not universal, it can be said to have evolved
to take different shapes in different species. The central
logic of theories of evolution of aging is that the force of
natural selection decreases with age of an individual [16, 17,