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PubMed2000

Shattered: Medawar's test tubes and their enduring legacy of chaos.

Medawar's 1952 paper 'An Unsolved Problem of Biology' underlies most subsequent theoretical work regarding the evolution of aging; it concludes that aging is accidental and could not have evolved; this prevents reconciling the growing body of evidence suggesting the existence of multiple, evolved, aging systems. The paper features a well-known thought experiment using test tubes to show why aging could not evolve. Medawar assumes that constant, random, breakage sufficiently represents lethal forces of nature; however, famine, drought, predation, disease, and accidents each uniquely affect populations. Predation is the only evolving force that continually invents new ways to kill members of the prey populations; thus all prey defenses to predation will eventually be defeated. Defenses to non-evolving or non-obligate lethal forces, however, should quickly evolve. Thus unevolving, identical test tubes cannot adequately represent biological populations. The example also ignores population booms and busts which often occur in nature. By ignoring these issues, Medawar examines only one population age distribution skewed towards younger individuals in predator-dominated environments while ignoring predator-free populations skewed towards older individuals after population crashes. Further, Medawar's test tubes lack meaningful competition for finite resources, and ignore declining fertility which occurs in all aging species. Medawar concludes that older individuals are too few in number to influence the population's gene pool for or against aging. This conclusion is found to be incorrect when variations in the age of reproductive senescence are introduced into a predator-free population.A new thought experiment with competing strains of algae corrects for these issues and shows that aging evolved and is retained so that groups retain enough genetic variability to allow for rapid evolution of a defense to novel predation. The example shows reasons why the rate of aging is directly linked to the reproductive rate, litter size, metabolic rate, reproductive senescence, and fixed body size. It also suggests that in the absence of predation, immortality would quickly evolve if not for the evolution of highly-conserved aging systems. Prior analysis of aging evolution is incorrect due to theorists' rejection of the idea of group selection. It is believed to be 'impossible' to select for mutations that are bad for the individual but good for the group. However, mutations that are neutral to young individuals which are only deleterious if expressed at older ages can accumulate in early-mortality, predator-dominated environments. Removing the predator allows deleterious mutation expression. Positive group selection then occurs amongst traits that are negative to the individual. Further, group selection is a universal force that occurs between local, non-breeding groups and not, as theorists propose, between distant groups of potentially interbreeding species. Local survivors migrate to replace extinct, related species. The antagonistic pleiotropy theory, which was created to salvage the idea of accidental aging, is examined and shown to be untenable. The hypothetical antagonistic pleiotropy genes that are beneficial to young while detrimental to old individuals, predicted to exist in the 1950s, are unlikely to exist, have not, and likely will not be found in sufficient quantity to participate in the aging process.

Aging↗

PubMedDate not supplied

Carnivoran paleoguilds of Africa: implications for hominid food procurement strategies.

Tool-using hominids, as carnivorous animals, would have been part of the various carnivore guilds present in Plio-Pleistocene Africa. Hominid dietary strategies must be understood within the larger context of carnivore behavior and ecology, as carnivorans could have affected the abilities of hominids to procure meat and/or marrow. The functional anatomy of extant and fossil carnivorans was examined to infer behaviors in fossil carnivorans that would have impacted on hominid dietary strategies in terms of carcass availability. Comparisons of guild structure were carried out to examine changes in carnivoran interactions and their implications for hominid behavior. Plio-Pleistocene carnivorans engaged in a wider range of behaviors than modern carnivorans. The sabertoothed felids Dinofelis and Megantereon probably did not provide much larger carcasses than modern species. Another sabertooth, Homotherium generated larger carcasses, but may have disarticulated and transported these carcasses. Fossil representatives of modern taxa may not have been equivalent ecologically within the carnivoran guild. Overall, hominids in eastern Africa probably had a greater range of scavenging opportunities than did those of southern Africa during the Plio-Pleistocene. Local and continent-wide extinction events in large-bodied carnivoran guilds from 1 to 2 Ma had a substantial effect on carcass availability and the risk to hominid scavengers. These structural changes in the carnivore guild may have provided an opportunity for hominids to widen their niche with respect to dietary behavior.

Africa↗

PubMed1983

Parasites at the origin of life.

This paper is concerned with parasitic virus-like particles and their hosts. It is proposed that parasitism must have occurred at an early stage of evolution, soon after the first self-reproducing systems had formed. When chemical building blocks for self-reproducing systems became scarce, current theories envision that some self-reproducing systems evolved the capability to synthesize materials for self-replication from chemical precursors in the environment. It is proposed that at about the same time parasitic systems (phages) arose that replicated at the expense of host systems by diverting host materials to the replication of their own genomes. With the aid of a mathematical model we demonstrate that host and phages can coexist in a stable equilibrium, depending upon the carrying capacity of the environment. If the latter falls below a threshold, then the parasites die out. A parasite that has the capability to integrate into the host genome is replicated along with it and thus escapes extinction during periods of population bottlenecks of the host population. The presence of phages creates evolutionary pressures favoring host defenses against them. Thus, modern bacteria are able to degrade most invading DNA (through restriction enzymes). Defense capabilities require a share of the genome, thus adding to the genetic complexity of organisms.

Biological Evolution↗

PubMed2002

The universal properties of stem cells as pinpointed by a simple discrete model.

The ability of a few stem-cells to repopulate a severely damaged bone marrow (BM) guarantees the stability of our physical existence, and facilitates successful BM transplantations. What are the basic properties of stem cells that enable the maintenance of the system's homeostasis? In the present work we attempt to answer this question by investigating a discrete (in time and phase-space) dynamical system. The model we present is shown to retrieve the essential properties of homeostasis, as reflected in BM functioning, namely, (a) to produce a constant amount of mature cells, and (b) to be capable of returning to this production after very large perturbations. The mechanism guaranteeing the fulfillment of these properties is extrinsic--negative feedback control in the micro-environment--and does not need additional stochastic assumptions. Nevertheless, the existence of a simple intrinsic control mechanism, a clock which determines the switch to differentiation, ascertains that the system does not admit non-trivial extinction states. This result may help clarifying some of the controversy about extrinsic versus intrinsic control over stem cell fate. It should be stressed that all conclusions are valid for any system containing progenitor and maturing cells.

Bone Marrow↗

PubMed2005

Fertility control in wildlife: humans as a model.

In a time of accelerated extinction rates and biodiversity loss, it may seem illogical to be concerned with methods limiting the reproduction of wild populations; however, there is an urgent need to inhibit the proliferation of a wide variety of species. The range of animals for which fertility control is desired makes the development of a single method impossible. The various reproductive strategies used by individual species, the desired outcome of contraceptive programs (reversible or irreversible; male or female directed) and our ignorance of the reproductive biology of many endangered species necessitate thorough species-specific investigations. As fertility control in humans is a reality and research into methods of fertility control is more advanced, humans serve as a model for developing contraceptive approaches for wild species. Population control by traditional methods (indirect or direct intervention by culling, poisoning, translocation, etc.) is increasingly unacceptable to the public, making human studies even more valuable for finding solutions to overabundant wild populations. This review compares and contrasts the range of contraceptive methods used in both wildlife and humans.

Animals↗

PubMed2004

Harvesting in a resource dependent age structured Leslie type population model.

We analyse the effect of harvesting in a resource dependent age structured population model, deriving the conditions for the existence of a stable steady state as a function of fertility coefficients, harvesting mortality and carrying capacity of the resources. Under the effect of proportional harvest, we give a sufficient condition for a population to extinguish, and we show that the magnitude of proportional harvest depends on the resources available to the population. We show that the harvesting yield can be periodic, quasi-periodic or chaotic, depending on the dynamics of the harvested population. For populations with large fertility numbers, small harvesting mortality leads to abrupt extinction, but larger harvesting mortality leads to controlled population numbers by avoiding over consumption of resources. Harvesting can be a strategy in order to stabilise periodic or quasi-periodic oscillations in the number of individuals of a population.

Age Distribution↗

PubMed1995

Quantitation of ultraviolet light-absorbing fractions of the cornea.

This study attempts to measure the quantitative contribution of major chemical fractions of the whole bovine cornea to ultraviolet (UV) absorption between 240 and 300 nm, with special attention to the biologically significant range of 290-300 nm. The cornea was divided into water-insoluble, nonprotein small water-soluble, water-soluble protein, and lipid-soluble fractions. The insoluble fraction (largely collagen) was solubilized by enzymatic digestion. Solutions of individual fractions equivalent to a constant mass of fresh cornea were scanned for absorption from 240 to 300 nm. The sum of the absorbances of the individual fractions closely approximated the absorbance of whole corneas at all wavelengths examined. The extinction coefficients of the lipid and water-soluble fractions were several times greater than that of the insoluble fraction throughout the studied spectrum. Yet, because of its large mass (75% of cornea dry weight), the insoluble fraction accounted for 40-50% of UV absorbance between 240 and 280 nm. However, in the range of 290-300 nm, the water-soluble plus lipid-soluble fractions accounted for 60-65% of the total absorption, with the water-soluble proteins alone accounting for 40-45% of the total. The soluble proteins comprised only approximately 17% of the cornea's dry weight. The special contribution of the water-soluble proteins to absorption was attributed to their relatively high tryptophan content (approximately 1.6% by weight). A 54-kDa protein, identified by others as tryptophan-rich, tumor-associated aldehyde dehydrogenase, accounted for approximately 30% of the total soluble protein mass.(ABSTRACT TRUNCATED AT 250 WORDS)

Absorption↗

PubMed2003

Gene flow and the coevolution of parasite range.

The geographic range of many parasites is restricted relative to that of their hosts. We study possible evolutionary mechanisms for this observation using a simple model that couples coevolution and demography. The model assumes that the environment consists of two habitats connected by movement and that coevolution is governed by quantitative traits. Our results demonstrate that host gene flow is an important determinant of parasite geographic range. Fluctuations in the rate of host gene flow cause shifts in parasite population densities and associated range expansions or contractions. In extreme cases, changing the rate of host gene flow can lead to global extinction of the parasite. Through a process we term demographic compensation, these shifts in parasite density may occur with little or no change in parasite adaptation to the host. As a consequence, reciprocal adaptation between host and parasite can become uncoupled from the rate of host gene flow.

Adaptation, Biological↗

PubMed1989

Effectiveness of zooprophylaxis in malaria control: a theoretical inquiry, with a model for mosquito populations with two bloodmeal hosts.

A model for a vector mosquito population with two bloodmeal hosts (man and a domestic animal) was developed to study the influences of domestic animals on the frequency of mosquito bites on man and the endemicity of human malaria. The vector population model, including blood-feeding success in the adult stage (depending on host density and biting efficiency) and density-dependent regulation in the larval stage, was combined with the Ross-Macdonald malaria transmission model. Model analyses suggested that introduction of domestic animals easily fed upon by mosquitoes increases mosquito density and, in some situations, frequency of mosquito bites on man and the infection rate of malaria through increased success of blood-feeding. Extinction of malaria was predicted only when an extremely large number of easily accessible (as compared to man) domestic animals are introduced. Limitations in the concept of zooprophylaxis and problems of livestock management in malaria control are discussed.

Animals↗

PubMed1999

On the theory of behavioral mechanics.

The Theory of Behavioral Mechanics is the behavioral analogue of Newton's laws of motion, with the rate of responding in operant conditioning corresponding to physical velocity. In an earlier work, the basic relation between rate of responding and sessions under two FI schedules and over a range of commonly used session values had been shown to be a power function. Using that basic relation, functions for behavioral acceleration, mass, and momentum are derived here. Data from other laboratories also support the applicability of a power function to VI schedules. A particular numerical value is introduced here to be the standard reference value for the behavioral force under the VI-60-s schedule. This reference allows numerical values to be calculated for the behavioral mass and momentum of individual animals. A comparison of the numerical values of the momenta of two animals can be used to evaluate their relative resistances to change, e.g., to extinction, which is itself viewed as a continuously changing behavioral force being imposed on the animal. This overall numerical approach allows behavioral force-values to be assigned to various experimental conditions such as the evaluation of the behavioral force of a medication dosage.

Animals↗

PubMed2002

Tangled nature: a model of evolutionary ecology.

We discuss a simple model of co-evolution. In order to emphasize the effect of interaction between individuals, the entire population is subjected to the same physical environment. Species are emergent structures and extinction, origination and diversity are entirely a consequence of co-evolutionary interaction between individuals. For comparison, we consider both asexual and sexually reproducing populations. In either case, the system evolves through periods of hectic reorganization separated by periods of coherent stable coexistence.

Animals↗

PubMed2003

Coexistence in a competitive parasitoid-host system.

The main objective of this work is to determine the conditions for coexistence and competitive exclusion in a discrete model for a community of three species: a stage-structured host and two competing parasitoids sharing the same host developmental stage. Coexistence of the community of the species is found to depend on the host life history parameters in the first place, and on competitive ability and parasitoid efficiency in the second place. In particular, parasitoids equilibrium densities are defined by the size of the refuge. Extinction is expected with low growth rate and with low adult survival. Host life histories are also associated with oscillations in population density, and depending on the combination of host adult survival from one generation to the next and host growth rate, the minimum of fluctuations approaches zero, implying a higher potential risk of extinction because of stochastic factors. Our results suggest that equally reduced survival of parasitoids in hosts parasitized by both species determines extinction of the parasitoid with lower population density, in contrast to the case when both parasitoids benefit with 50% of all doubly parasitized hosts, leading to the hypothesis that a community where competitors in multiparasitized hosts die, easily becomes extinct. Competitive exclusion is expected for highly asymmetric competitive interactions, independent of population densities, allowing us to hypothesize that coexistence of competitors in systems with limited resources and refuges is associated with a clearly defined competitive hierarchy.

Animals↗

PubMed1999

Evolutionary dynamics of seed size and seedling competitive ability.

We present a model for the evolutionary dynamics of seed size when there is a trade-off between seed size and seed number, and seedlings from large seeds are better competitors and have a higher precompetitive survival than seedlings from small seeds. We find that strong competitive asymmetry, high resource levels, and intermediate harshness of the precompetitive environment favor coexistence of plants with different seed sizes. If the evolution of seed size is mutation-limited and single mutations have only a small phenotypic effect, then an initially monomorphic population reaches the final evolutionarily stable polymorphic state through one or more discrete evolutionary branching events. At each such branching event, a given lineage already present in the population divides into two phenotypically diverging daughter lines, each with its own seed size. If the precompetitive survival of seeds and seedlings is high for small and large seeds alike, however, evolutionary branching may be followed by the extinction of one or more lineages. Various results presented here are model-independent and point the way to a more general evolutionary bifurcation theory describing how the number and stability properties of evolutionary equilibria may change as a consequence of changes in model parameters.

Biological Evolution↗

PubMed1999

Population dynamics under parasitic sex ratio distortion.

We analyse the population dynamic effects of sex ratio distortion by vertically transmitted, feminizing parasites. We show that, for diploid hosts, sex ratio distortion may lead to extinction as males become too rare to maintain the host population through reproduction. Feminizers can magnify Allee effects, broadening the range of conditions leading to extinction of small populations. Depending on male mating constraints and strength of density dependence, feminizers may either increase or decrease the equilibrium host density. Under conditions leading to deterministic host extinction, stochastic elimination of the parasite may allow the host population to recover. Hence, infection by parasitic sex ratio distorters may be transient in finite populations. We consider the implications of this process for parasite prevalence, host population regulation, and sex ratio evolution.

Animals↗

PubMed1992

Liposome shrinkage and swelling under osmotic-diffusional stress: evaluation of kinetic parameters from spectrophotometric measurements.

Analyses of spectrophotometric measurements of shrinkage and swelling of liposomes under osmotic-diffusional stress tacitly assume an 'empirical' inverse relationship between the absorbance (A) and the liposomal volume (v). In this paper, we have proposed an alternate more explicit relationship: A = EcL e - alpha Lv/1-Lv where L is liposomal concentration, Ec the extinction coefficient and alpha a dimensionless parameter. The exponential term, in essence, defines the partitioning of aqueous volume into intra- and extra-liposomal compartments. Experimental data obtained with glycerol as model compound are used to test the validity and internal consistency of the proposed formalism.

Kinetics↗

PubMed2001

Ancient DNA.

DNA that has been recovered from archaeological and palaeontological remains makes it possible to go back in time and study the genetic relationships of extinct organisms to their contemporary relatives. This provides a new perspective on the evolution of organisms and DNA sequences. However, the field is fraught with technical pitfalls and needs stringent criteria to ensure the reliability of results, particularly when human remains are studied.

Animals↗

PubMed1994

Extinction rates can be estimated from molecular phylogenies.

Molecular phylogenies can be used to reject null models of the way we think evolution occurred, including patterns of lineage extinction. They can also be used to provide maximum likelihood estimates of parameters associated with lineage birth and death rates. We illustrate: (i) how molecular phylogenies provide information about the extent to which particular clades are likely to be under threat from extinction; (ii) how cursory analyses of molecular phylogenies can lead to incorrect conclusions about the evolutionary processes that have been at work; and (iii) how different evolutionary processes leave distinctive marks on the structure of reconstructed phylogenies.

Animals↗

PubMed2003

Testing for equal rates of cladogenesis in diverse taxa.

Taxa differ widely in numbers of species, which may be due either to chance alone or to factors that cause differences in speciation and extinction rates between taxa. To test whether an observed distribution of species over taxa differs from the distribution expected from chance alone, one must take into account that neither speciation nor extinction rates are known. This paper introduces a way to estimate speciation and extinction probabilities from the distribution of extant species over families and to test whether the observed distribution is different from expected. Application of this procedure to the distributions of bird, hexapod, primate, and angiosperm species over taxa provides statistical evidence of differences in rates of cladogenesis between taxa.

Animals↗