Living fast and dying young: A comparative analysis of life-history variation among mammals

Recent comparative studies point to the importance of mortality schedules as determinants in the evolution of life-history characteristics. In this paper, we compare patterns of mortality from natural populations of mammals with a variety of life histories. We find that, after removing the effects of body weight, mortality is the best predictor of variation in life-history traits. Mammals with high levels of natural mortality tend to mature early and give birth to small offspring in large litters after a short gestation, before and after body size effects are factored out. We examine the way in which life-history traits relate to juvenile mortality versus adult mortality and find that juvenile mortality is more highly correlated with life-history traits than is adult mortality. We discuss the necessity of distinguishing between extrinsic sources of mortality (e.g. predation) and mortality caused by intrinsic sources (e.g. costs of reproduction), and the role that ecology might play in the evolution of patterns of mortality and fecundity. We conclude that these results must be explained not simply in the light of the demographic necessity of balancing mortality and fecundity, but as a result of age-specific costs and benefits of reproduction and parental investment. Detailed comparative studies of mortality patterns in natural populations of mammals offer a promising avenue towards understanding the evolution of life-history strategies.     

Fast and slow axonal transport-different methodological approaches give complementary information: contributions of the stop-flow/crush approach

This minireview describes experiments in short term crush operated rat nerves, to study endogenous substances in anterograde and retrograde fast axonal transport. Immunofluorescence was used to recognize transported antigens, and cytofluorimetric scanning was employed to quantitate different antigens which had accumulated proximal and distal to the crushes. Vesicle membrane components p38 (synaptophysin) and SV2 accumulated on both sides of a crush. This was expected from a number of studies from different laboratories. Surface associated molecules, however, like synapsins and rab3a, have been studied by other groups with biochemical methods, and suggested to be transported with slow transport. The crush method, however, revealed that a considerable fraction of these two substances are transported with the fast transport system, and, thus, associated with fast transported organelles in the living neuron. Evidently, more than one technique is required to give a more complete picture of intraneuronal transport related events.Special issue dedicated to Dr. Sidney Ochs.     

Demographic stochasticity versus spatial variation in the competition between fast and slow dispersers

Dispersal is an important strategy that allows organisms to locate and exploit favorable habitats. The question arises: given competition in a spatially heterogeneous landscape, what is the optimal rate of dispersal? Continuous population models predict that a species with a lower dispersal rate always drives a competing species to extinction in the presence of spatial variation of resources. However, the introduction of intrinsic demographic stochasticity can reverse this conclusion. We present a simple model in which competition between the exploitation of resources and stochastic fluctuations leads to victory by either the faster or slower of two species depending on the environmental parameters. A simplified limiting case of the model, analyzed by closing the moment and correlation hierarchy, quantitatively predicts which species will win in the complete model under given parameters of spatial variation and average carrying capacity.     

Resource availability, mortality, and fertility: a path analytic approach to global life-history variation

Humans exhibit considerable diversity in timing and rate of reproduction. Life-history theory (LHT) suggests that ecological cues of resource richness and survival probabilities shape human phenotypes across populations. Populations experiencing high extrinsic mortality due to uncertainty in resources should exhibit faster life histories. Here we use a path analytic (PA) approach informed by LHT to model the multiple pathways between resources, mortality rates, and reproductive behavior in 191 countries. Resources that account for the most variance in population mortality rates are predicted to explain the most variance in total fertility rates. Results indicate that resources (e.g., calories, sanitation, education, and health-care expenditures) influence fertility rates in paths through communicable and noncommnunicable diseases. Paths acting through communicable disease are more strongly associated with fertility than are paths through noncommunicable diseases. These results suggest that a PA approach may help disaggregate extrinsic and intrinsic mortality factors in cross-cultural analyses. Such knowledge may be useful in developing targeted policies to decrease teenage pregnancy, total fertility rates, and thus issues associated with overpopulation.     

Relationships of latitude,altitude, and body size to litter size and mean annual production of offspring inPeromyscus

Litter size was positively correlated with latitude and altitude but not with production of offspring or with body size in Peromyscus. Increased litter size in northern populations probably reflects shortening the breeding season by climate and not a greater mortality rate at northern latitudes compared to southern latitudes. Production of offspring was negatively correlated with body size but not with latitude, altitude, or litter size. This is probably due to larger species living longer and taking longer to mature.     

Early life-history dynamics of Caribbean coral species on artificial substratum: the importance of competition, growth and variation in life-history strategy

The development of a coral community was monitored for 6 years (1998–2004) on 46 m² of artificial settlement substrate in Curaçao, Netherlands Antilles. Growth and survival of recruits (n=1385) belonging to 16 different species were quantified in relation to characteristics of the benthic community developing around them. The early life history dynamics (i.e. growth rate, growth strategy and survival) of corals differed among species although these differences were small for species occupying similar habitats (i.e. underside versus topside of substratum). In contrast to recruit survival, juvenile growth rates were highly variable and unrelated to benthic community structure, at least at the scale of this study. Competing benthic organisms affected coral recruitment success through space preemption (mainly by macroalgae) or recruit overgrowth (mainly by sponges). The results highlight the small spatial scale (mm–cm) at which the processes responsible for recruitment success or failure occur and emphasize the need to include such small-scale observations in studies of coral early life-phase dynamics.     

Energy budgets, growth rates, and thermal constraints: toward an integrative approach to the study of life-history variation

Variation in thermal constraints on activity has been hypothesized to be an important ecological source of geographic variation in growth rates of juvenile eastern fence lizards Sceloporus undulatus. However, most of the evidence to support this hypothesis is either inferential or indirect. In this study, I quantitatively compared thermal constraints on activity and their relationship to growth rates of free-ranging juvenile fence lizards from two extremes of the range of variation in growth rate (Nebraska and New Jersey) used in a reciprocal transplant experiment. I also examined energy allocation made to growth and storage by yearling lizards. Reduced growth rates in New Jersey of normally fast-growing hatchlings from Nebraska were associated with a more stringent thermal constraint on activity corresponding to a 2-3-h shorter predicted daily activity period in New Jersey compared to Nebraska. The thermal constraint on activity was particularly strong (24% less time available in New Jersey compared to Nebraska) during the period when hatchling lizards emerge (August-October). An 8% reduction in total activity time available over the course of a single year was associated with a 7% reduction in the total amount of energy accumulated by lizards in New Jersey. Differences in the total amount of energy available for allocation were also accompanied by differences in how energy was allocated. Lizards from New Jersey had an allocatable energy pool of approximately 40.34 kJ (88% to growth, 12% to storage, and 0% to reproduction). Lizards from Nebraska had an allocatable pool of 43.44 kJ (22% to growth, 18% to storage, and 60% to reproduction). This study joins others in advocating and illustrating an integrative approach to determining the causes and consequences of life-history variation by combining experimental, comparative, and phylogenetic methods in a single system.     

Breaking cover: neural responses to slow and fast camouflage-breaking motion

Primates need to detect and recognize camouflaged animals in natural environments. Camouflage-breaking movements are often the only visual cue available to accomplish this. Specifically, sudden movements are often detected before full recognition of the camouflaged animal is made, suggesting that initial processing of motion precedes the recognition of motion-defined contours or shapes. What are the neuronal mechanisms underlying this initial processing of camouflaged motion in the primate visual brain? We investigated this question using intrinsic-signal optical imaging of macaque V1, V2 and V4, along with computer simulations of the neural population responses. We found that camouflaged motion at low speed was processed as a direction signal by both direction- and orientation-selective neurons, whereas at high-speed camouflaged motion was encoded as a motion-streak signal primarily by orientation-selective neurons. No population responses were found to be invariant to the camouflage contours. These results suggest that the initial processing of camouflaged motion at low and high speeds is encoded as direction and motion-streak signals in primate early visual cortices. These processes are consistent with a spatio-temporal filter mechanism that provides for fast processing of motion signals, prior to full recognition of camouflage-breaking animals.     

Acetylcholinesterase molecular forms in the fast or slow muscles of the chicken and the pigeon

Molecular forms of acetylcholinesterase (AChE) were examined in various skeletal muscles of the chicken and the pigeon. In chicken pectoralis m., AChE was found to be restricted to endplate containing segments, and no asymmetric form could be detected in aneural samples. In the chicken muscles studied, a relation has been established between globular (G1,G2,G4) forms or asymmetric (A8,A12) forms, and muscle fibre types. Asymmetric forms are preponderant in fast-twitch muscles, whereas in slow tonic muscles 80% of the AChE activity is due to globular forms. However, comparison with pigeon muscles shows that AChE chicken muscle patterns may not be generalized.     

Structure and diversity of stream invertebrate assemblages: the influence of temperature with altitude and latitude

1. Structure and diversity of the macroinvertebrate fauna were studied in relation to altitude and latitude among three groups of streams from Ecuador (lowland: 100–600 m, Central Valley: 2600–3100 m, páramo: 3500–4000 m), and one group from the temperate lowland region of Denmark. The streams in the four regions were comparable with regard to physical characteristics such as size, current and substratum.
2. In terms of faunal composition the Ecuadorian highland streams bore more resemblance to the Danish lowland streams than the Ecuadorian lowland streams. The greater similarity between the Ecuadorian highland and the Danish streams, however, was due to the large number of insect families in the Ecuadorian lowlands, many of which were not found in the other regions. Of ten physico-chemical parameters measured, maximum stream temperature explained by far the most variability in faunal composition.
3. The number of insect orders and families increased linearly with maximum stream temperature and therefore decreased with altitude and latitude. A compilation of literature data on insect richness and maximum water temperature from streams around the world confirmed this pattern, yielding a common linear relation for both temperate and tropical streams. This pattern may arise due to a direct temperature effect on speciation but is probably also related to geological history and the influence of climatic changes on stream ecosystems. We estimate that small, tropical, lowland streams have, on average, a two- to fourfold higher species richness than temperate lowland streams.     

Growth rate plasticity to temperature in two damselfly species differing in latitude: contributions of behaviour and physiology

Plasticity in growth rate may be driven by behavioural and physiological mechanisms. Although these underlying mechanisms have direct implications for the importance of ecological and physiological costs associated with rapid growth, the contribution of behaviour and physiology to temperature-mediated plasticity in growth rate has largely been neglected. We studied the temperature-dependence of growth rate and its underlying behavioural and physiological mechanisms in two congeneric damselfly species that differ in latitudinal distribution. Larvae were reared from the egg stage at three temperatures (17°C, 22°C and 27°C). Within each species, growth rates showed a quadratic response curve with an optimum at 22°C. Behaviour, as measured by food intake, and physiology, as measured by growth efficiency and heartbeat as proxy for metabolic rate, jointly contributed to this temperature-induced plasticity in growth rate. At each temperature, growth rates were higher in the northern species. In line with the few other studies that compared northern and southern populations, both an increased food uptake and growth efficiency caused this pattern. Together with previous studies that focused on the population level, our results tentatively suggest that not only the latitudinal patterns in growth rate but also the mechanistic basis are similar at the species and at the population level.     

Analysis of genetic variation affecting the relative activities of fast and slow ADH dimers in Drosophila melanogaster heterozygotes

When Adh^F/Adh^S heterozygote homogenates are stained after electrophoresis, considerable variation is observed in the activity ratio of the FF dimer to the SS dimer. Two Adh^S strains showed a sharp, consistent difference when crossed to a common Adh^F strain. Optical scanning and genetic analysis confirmed that this difference originates close to the Adh locus. Since the morphs varied concordantly in their activities on numerous alcohols, and since aging and heat-treatment experiments failed to reveal a stability difference, it is proposed that the difference is regulatory in nature, affecting ADH synthesis and primarily cis-acting. A survey of wild flies revealed additional variation in the FF/SS activity ratio. Further genetic analysis showed that the basis of this variation is not restricted to the second chromosome. Furthermore, modification of the activity ratio implies some degree of allelespecificity on the part of the modifiers.This work was supported in part by money collected by Jewish Community of Iowa City, Iowa, and by NSF Grant 76-01903 to Roger Milkman.     

Ecogeographic analysis of morphological and life-history variation in the Italian treefrog

We describe the spatial pattern of variation in body size, body shape, and adult population age structure of the Italian treefrog (Hyla intermedia). By means of a non-parametric test of matrix association (the Partial Mantel Test), we investigate the role of climate and geography in explaining the patterns of phenotypic variation. Body size is the most important source of morphometric variation, both at intra- and inter-populational levels. Body shape, despite its secondary role in explaining variation, still maintains high discriminative power among populations. Age structure differs significantly among populations, since body size correlates positively to age, these differences partly explain geographic variation of body size. The patterns of variation in both age and body size are congruent with climate variation, suggesting causal relationship. However, we find no evidence for adaptive explanations and suggest that climate-induced demographically differential survivorship might be the proximate cause responsible for the associations observed. Unlike body size, body shape varies congruently with the pattern of between-population geographic distances, which we show to be highly correlated with their genetic distances, thus suggesting isolation by distance as the proximate cause of the association.     

Regenerated rat fast muscle transplanted to the slow muscle bed and innervated by the slow nerve,exhibits an identical myosin heavy chain repertoire to that of the slow muscle

 The hypothesis that the limited adaptive range observed in fast rat muscles in regard to expression of the slow myosin is due to intrinsic properties of their myogenic stem cells was tested by examining myosin heavy chain (MHC) expression in regenerated rat extensor digitorum longus (EDL) and soleus (SOL) muscles. The muscles were injured by bupivacaine, transplanted to the SOL muscle bed and innervated by the SOL nerve. Three months later, muscle fibre types were determined. MHC expression in muscle fibres was demonstrated immunohistochemically and analysed by SDS-glycerol gel electrophoresis. Regenerated EDL transplants became very similar to the control SOL muscles and indistinguishable from the SOL transplants. Slow type 1 fibres predominated and the slow MHC-1 isoform was present in more than 90% of all muscle fibres. It contributed more than 80% of total MHC content in the EDL transplants. About 7% of fibres exhibited MHC-2a and about 7% of fibres coexpressed MHC-1 and MHC-2a. MHC-2x/d contributed about 5–10% of the whole MHCs in regenerated EDL and SOL transplants. The restricted adaptive range of adult rat EDL muscle in regard to the synthesis of MHC-1 is not rooted in muscle progenitor cells; it is probably due to an irreversible maturation-related change switching off the gene for the slow MHC isoform. Accepted: 11 June 1996     

Reaction of formate with the fast form of cytochrome oxidase: a model for the fast to slow conversion.

The ability to isolate preparations of cytochrome oxidase which are highly homogeneous has facilitated a study of the effects of various reagents on the purified enzyme. The addition of either sodium formate, formamide, formaldehyde, or sodium nitrite to enzyme which reacts in a single rapid kinetic phase with cyanide causes a blue-shift of 4-6 nm of the net (cytochrome a + cytochrome a3) Soret maximum. Only the derivative prepared by adding sodium formate demonstrates measurable intensity in the g' = 12 region of the low-temperature electron paramagnetic resonance (EPR) spectrum. This g' = 12 resonance is characteristic of cytochrome oxidase which has undergone a modification at the binuclear center and thereby reacts sluggishly with cyanide. As the site of cyanide binding in resting enzyme as been demonstrated to be CuB [Yoshikawa, S., & Caughey, W.S. (1990) J. Biol. Chem. 265, 7945-7958], it is proposed that formate can bind to CuB and the fast to slow transition is rationalized by using this proposal. The g' = 12 signal is also produced upon the addition of sodium formate to mitochondrial preparations, suggesting that the species responsible for this behavior may have possible physiological relevance. Physical properties of the formate derivative and data for other reagents reacted with the fast-reacting enzyme preparation are presented.     

High levels of liver antioxidants are associated with life-history strategies characteristic of slow growth and high survival rates in birds

Antioxidants have a large potential to coevolve with life-histories because of their capacity to counteract the negative effects of free radicals on fitness. However, only a few studies have explored the association between antioxidant levels and life-history strategies comparing a large number of species. Here we used an extensive dataset of 125 species of birds to investigate the association between concentrations of antioxidants (carotenoids and vitamin E) in the liver, which is the main storage organ for fat-soluble antioxidants, and life-history and morphology. We found that high liver antioxidant concentrations were associated with life-history strategies characterized by "live slow, die old", in clear contrast to previous studies reporting a relationship between high plasma antioxidants and life-histories characterized by "live fast, die young". Thus, high carotenoid concentrations were present in species with large body, brain and egg sizes, high absolute metabolic rate and a resident lifestyle, while high vitamin E concentrations were present in species with large brain size and long life span and incubation period. Our results indicate that antioxidants and life-histories coevolve, and that this may be mediated by positive fitness consequences of the accumulation of liver antioxidants, as species with higher antioxidant levels live longer.     

On the use of metaphor to understand, explain, or rationalize redundant genes in yeast

The proposal that yeast, and cells in general, contains redundant genes that enable cells to survive mutational change has been supported by experiments and a strong metaphor. The redundant gene proposal is analyzed, and it is noted that there are many problems with the redundant gene model. An alternative metaphor is suggested to explain the genetic composition of a yeast culture.