Air flow and convective heat loss from small mammals in laboratory metabolism chambers

1. 1.Thermal conductance was determined from cooling curves of Mus domesticus carcasses at air flow rates ranging from still air to 0.91·min⁻¹ STP. Thermal conductance was constant over this range of air flows. This indicates that free convection predominates over forced convection at these air flow rates.

2. 2.While non single set of conditions will be appropriate for all experiments, free convection conditions are appropriate for determination of minimal thermal conductance.

Effect of wind speed on loss of water from Nostoc flagelliforme colonies

The effects of wind speed on loss of water from N. flagelliforme colonies were investigated indoors in an attempt to assess its ecological significance in field. Wind enhanced the process of waterloss; the half-time of desiccation at wind speeds of 2.0 and 3.4 m s^-1 was, respectively, shortened to one-third and one-fifth at 20°C and, to one-sixth and one-eighth at 27°C that of still air. Photosynthetic efficiency was not affected before the wet alga lost about 50% water. This revised version was published online in June 2006 with corrections to the Cover Date.     

Body mass, population density, and offspring number in mammals

The negative relationship between population density and body mass with the body mass exponent of -0.75 implies that the energy flow through populations of small- and large-bodied species is the same, for individual metabolism scales to body mass raised to the power of +0.75. This relationship called the energetic equivalence rule, has often been observed for mammal species assemblages studied at regional scales. Here we suggest a demography-based mechanism that may generate it. Having analyzed about 130 literature sources, mostly in Russian, we collected demography and body-mass data for 88 mammalian species from the territory and coastal waters of the former Soviet Union. The data were used to construct a number of interspecific relationships. It is shown that (1) the number of offspring per lifetime is approximately inversely proportional to the relative mass at birth (the exponent is not significantly different from -1), (2) the average lifespan is proportional to body mass to the 0.25 power, (3) body mass at birth is proportional to the adult body mass. We develop a simple theory to demonstrate that relations (1) to (3) entail the energetic equivalence rule. The theory also allows us to explain violation of this rule (in non-flying birds, for example), namely, to predict the exponent of relation (1) for any given exponent of the relation between population density and body mass. This is possible because relations (2) and (3) are likely to more universally hold than relation (1). Finally, since natural selection acts on individual traits rather than on population-level ones such as population density, the theory opens up the way to an evolutionary explanation for the energetic equivalence rule.     

Estimating metabolic heat loss in birds and mammals by combining infrared thermography with biophysical modelling

Infrared thermography (IRT) is a technique that determines surface temperature based on physical laws of radiative transfer. Thermal imaging cameras have been used since the 1960s to determine the surface temperature patterns of a wide range of birds and mammals and how species regulate their surface temperature in response to different environmental conditions. As a large proportion of metabolic energy is transferred from the body to the environment as heat, biophysical models have been formulated to determine metabolic heat loss. These models are based on heat transfer equations for radiation, convection, conduction and evaporation and therefore surface temperature recorded by IRT can be used to calculate heat loss from different body regions. This approach has successfully demonstrated that in birds and mammals heat loss is regulated from poorly insulated regions of the body which are seen to be thermal windows for the dissipation of body heat. Rather than absolute measurement of metabolic heat loss, IRT and biophysical models have been most useful in estimating the relative heat loss from different body regions. Further calibration studies will improve the accuracy of models but the strength of this approach is that it is a non-invasive method of measuring the relative energy cost of an animal in response to different environments, behaviours and physiological states. It is likely that the increasing availability and portability of thermal imaging systems will lead to many new insights into the thermal physiology of endotherms.     

Flight speed of seabirds in relation to wind speed and direction

We studied flight speed among all major seabird taxa. Our objectives were to provide further insight into dynamics of seabird flight and to develop allometric equations relating ground speed to wind speed and direction for use in adjusting seabird density estimates (calculated from surveys at sea) for the effect of bird movement. We used triangulation at sea to estimate ground speeds of 1562 individuals of 98 species. Species sorted into 25 “groups” based on similarity in ground speeds and taxonomy. After they were controlled for differences inground speed, the 25 groups sorted into eight major “types” on the basis of response to wind speed and wind direction. Wind speed and direction explained 1664% of the variation in ground speed among seabird types. For analyses on air speed (ground speed minus apparent wind speed), we divided the 25 groups according to four flight styles: gliding, flap-gliding, glide-flapping and flapping. Tailwind speed had little effect on air speed of gliders (albatrosses and large gadfly petrels), but species that more often used flapping decreased air speed with increase in tailwinds. All species increased air speeds significantly with increased headwinds. Gliders showed the greatest increase relative to increase in headwind speed and flappers the least. With tailwind flight, air speeds were greatest among species with highest wing loading for each flight style except gliders, which showed no relationship. For headwind flight, species with higher wing loading had higher air speeds; however, the relation was weaker in flappers compared with species using some amount of gliding. In contrast, analyses for air speed ratio (i.e. difference between air speed in acrosswinds [with no apparent wind] and speed flown into headwinds, or with tailwinds, divided by speed acrosswind) revealed that among species using some flapping, and with lower wing loading (surface-feeding shearwaters, small gadfly petrels, storm petrels, phalaropes, gulls and terns), adjusted air speeds more than those with higher wing loading (alcids, “diving shearwaters”, “Manx-type shearwaters”, pelicans, boobies and cormorants). As a result, most flappers of low wing loading flew much faster than V_mr (the most energy efficient air speed per distance flown) when flying into headwinds. We suggest that better-than-predicted gliding performance with acrosswinds and tailwinds of large gadfly petrels, compared with albatrosses, resulted from a different type of “soaring” not previously described in seabirds.     

Locomotor mode, maximum running speed, and basal metabolic rate in placental mammals

The locomotor performance (absolute maximum running speed [MRS]) of 120 mammals was analyzed for four different locomotor modes (plantigrade, digitigrade, unguligrade, and lagomorph-like) in terms of body size and basal metabolic rate (BMR). Analyses of conventional species data showed that the MRS of plantigrade and digitigrade mammals and lagomorphs increases with body mass, whereas that of unguligrade mammals decreases with body mass. These trends were confirmed in plantigrade mammals and lagomorphs using phylogenetically independent contrasts. Multiple regression analyses of MRS contrasts (dependent variable) as a function of body mass and BMR contrasts (predictor variables) revealed that BMR was a significant predictor of MRS in the complete data set, as well as in plantigrade and nonplantigrade mammals. However, there was severe multicollinearity in the nonplantigrade model that may influence the interpretation of these models. Although these data show mass-independent correlation between BMR and MRS, they are not necessarily indicative of a cause-effect relationship. However, the analyses do identify a negligible role of body size associated with MRS once phylogenetic and BMR effects are controlled, suggesting that the body size increase in large mammals over time (i.e., Cope's rule) can probably rule out MRS as a driving variable.     

Air movement and heat loss from sheep. II. Thermal insulation of fleece in wind

Penetration of an animal's coat by wind reduces its thermal insulation and increases heat loss to the environment. From studies of the sensible heat loss from a life-sized model sheep covered with fleece, the average fleece resistance -rf(s cm-1) was related to windspeed u (m s-1) by 1/-rf(u) = 1/-rf(0)+cu, where c is a dimensionless constant. As c is expected to be inversely proportional to coat depth l, the more general relation -k(u) = -k(0)+c'u was evaluated, where -k = l/-rf is the thermal diffusivity (cm2 s-1) of the fleece and c' = cl is another constant (cm). The orientation of the model to the wind had little effect on the bulk resistance of the fleece, but the resistance on the windward side was substantially lower than on the leeward side.     

Solar heat gain in a desert rodent: unexpected increases with wind speed and implications for estimating the heat balance of free-living animals

We quantified metabolic power consumption as a function of wind speed in the presence and absence of simulated solar radiation in rock squirrels, Spermophilus variegatus, a diurnal rodent inhabiting arid regions of Mexico and the western United States. In the absence of solar radiation, metabolic rate increased 2.2-fold as wind speed increased from 0.25 to 4.0 m·s^-1. Whole-body thermal resistance declined 56% as wind speed increased over this range, indicating that body insulation in this species is much more sensitive to wind disruption than in other mammals. In the presence of 950 W·m^-2 simulated solar radiation, metabolic rate increased 2.3-fold as wind speed was elevated from 0.25 to 4.0 m·s^-1. Solar heat gain, calculated as the reduction in metabolic heat production associated with the addition of solar radiation, increased with wind speed from 1.26 mW·g^-1 at 0.25 m·s^-1 to 2.92 mW·g^-1 at 4.0 m·s^-1. This increase is opposite to theoretical expectations. Both the unexpected increase in solar heat gain at elevated wind speeds and the large-scale reduction of coat insulation suggests that assumptions often used in heat-transfer analyses of animals can produce important errors.Abbreviations absorptivity of coat to solar radiation - kinematic viscosity of air (mm²·s^-1) - reflectivity of coat to solar radiation - a r _B expected at zero wind speed (s·m^-1) - A _P projected surface area of animal on plane perpendicular to solar beam (cm²) - A _SKIN skin surface area (cm²) - b Coefficient describing change in r _B with change in square-root of wind speed (s^1.5·m^1.5) - d hair diameter (m) - d characteristic dimension of animal (m) - D _H thermal diffusivity of air (m²·s^-1) - E evaporative heat loss (W·m^-2) - I probability per unit coat depth that photon will strike hair - k constant equalling 1200 J·m^-3·°C^-1 - l _C coat depth m) - l _H hair length (m) - M metabolic rate (W·m^-2) - n density of hairs of skin (m^-2) - Q _A solar heat gain to animal (W·m^-2) - Q _I solar irradiance intercepted by animal (W·m^-2) - RQ respiratory quotient - r _A thermal resistance of boundary layer (s·m^-1) - r _B whole-body thermal resistance (s·m^-1) - r _E thermal resistance between animal surface and environment s·m^-1) - r _R radiative resistance (s·m^-1) - r _S sum of r _B and r _E at 0.25 m·s^-1 (s·m^-1) - r _T tissue thermal resistance s·m^-1) - T _AIR air temperature (°C) - T _B body temperature (°C) - T _E operative temperature of environment (°C) - T _ES standard operative temperature of environment (°C) - u wind speed (m·s^-1)     

Responses to heat and cold in lower mammals

A survey has been made of the literature pertaining to temperature regulation in the lower mammals.The following scheme represent a tentative outline of one probable evolutionary sequence of homeothermy: the first successful efforts to maintain a fairly uniform level of body temperature began on the psycho-physiological or behavioural level. The first directly physiological factor in the maintenance of homeothermy seems to have been a capacity for variation in metabolic heat production.The monotremes provides an example of a temperature regulation based solely on behaviour patterns and variation in heat production. The next factor to appear in the gradual development of refined homeothermy has been the regulation of physical heat exchange. A gradual improvement of such function is discernible along the phylum also correlated with increased thermal stress imposed by the environment. Regulation of physical heat exchange is first generally apparent in the marsupial order. The thermostatic arrangements responsible for efficient body temperature regulation have been developed gradually becoming increasingly complex and accurate. It is discussed how an efficient central nervous thermostatic control seems to have been the last factor developed to perfection in homeothermy.

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Zusammenfassung übersichtsrefrat über die Temperaturregulation in niederen SÄugern. Das folgende Schema ist ein vorlÄufiger Entwurf über die wahrscheinliche Ausbildung der Homeothermie wÄhrend der Evolution:die ersten wirksamen Einrichtungen zur Erhaltung einer ziemlich uniformen Korpertemperatur waren psycho-physiologische Funktionen und Eigenarten im Verhalten. Die erste ausschliesslich physiologische Funktion zur Erhaltung der Homeothermie scheint die FÄhigkeit zur Änderung der WÄrmebildung im Stoffwechsel gewesen zu sein.Die Monotremata sind ein Beispiel für Temperaturregulation, die allein durch Verhaltensformen und Änderung der WÄrmebildung bestimmt wird. Der nÄchste Schritt wÄhrend der graduellen Entwicklung zur ausgebildeten Homeothermie war die Regulierung des physikalischen WÄrmeaustausches.Eine zunehmende Verbesserung dieser Funktionen ist in dem Stamm zu beobachten. Sie ist verbunden mit verstÄrkter Hitzebelastung durch die Umwelt.Die Regulierung des physikalischen WÄrmeaustausches ist bei den Marsupialia erstmalig nachweisbar.Die Einrichtungen für wirksame Körpertemperaturregulation haben sich langsam entwickelt und wurden immer komplexer und genauer. Die Körpertemperaturkontrolle durch das Zentralnervensystem scheint die letzte Einrichtung zur Vervollkommung der Homeothermie gewesen zu sein.

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Résumé Ce travail passe en revue la régulation thermique des petits mammifères.Le schéma provisoire suivant indique les mécanismes possibles d'apparition de l'homéothermie pendant l'évolution: les premiers dispositifs efficaces maintenant une température corporelle à peu près constante étaient des fonctions psycho-physiologiques et des particularités du comportement. La première fonction purement physiologique contribuant à l'homéothermie semble avoir été la faculté de modifier la production de chaleur métabolique. Les monotrèmes sont un exemple de régulation thermique produite uniquement par des formes de comportement et par des changements de la production de chaleur. Le développement de l'homéothermie complète progressa ensuite par la régulation des échanges physiques de chaleur.Le perfectionnement de ces fonctions s'observe chez le phylum.Elle est liée à une surcharge de chaleur en provenance du milieu externe. La régulation des échanges physiques de chaleur est atteinte en premier par les marsupiaux. Les dispositifs efficaces de régulation de la température corporelle se sont développés lentement et sont devenus toujours plus précis et plus complexes. Le contrÔle de la température corporelle par le système nerveux central semble Être le dispositif le plus récent de l'homéothermie complète.

     

The effect of wind on the rate of heat loss from avian cup-shaped nests

Forced convection can significantly influence the heat loss from birds and their offspring but effects may be reduced by using sheltered micro-sites such as cavities or constructing nests. The structural and thermal properties of the nests of two species, the spiny-cheeked honeyeater (Acanthagenys rufogularis) and yellow-throated miner (Manorina flavigula), were measured in relation to three wind speeds. Nest dimensions differ between the two species, despite the similar body mass of the incubating adults, however nest conductance is comparable. As wind speed increases, so does the rate of heat loss from the nests of both species, and further still during incubation recesses. The significance of forced convection through the nest is a near-doubling in heat production required by the parent, even when incubating at relatively low wind speeds. This provides confirmation that selecting a sheltered nest site is important for avian reproductive success.     

Tree density,pollen-ovule ratio,and sex ratio in dioecious wind pollinated trees

The comparative reproductive success of male and female trees of a wind pollinated species is studied by simulation. Young trees are recruited into an established forest. When the sex ratio of the established forest is even, the reproductive success of both male and female recruits increases with forest density, and in general a recruit of either sex contributes the same number of successful gametes. When the sex ratio of the established forest deviates from unity, the recruit of the minority sex has a reproductive advantage over the recruit of the majority sex. Because a forest with a biased sex ratio can produce the same level of pollination as a forest with an even sex ratio, the quantity of pollen reaching the stigma of a flower does not convey enough information to permit the sex ratio of progeny to be biased adaptively as has been claimed. The male recruit has a slight advantage over the female recruit when the pollen-ovule ratio is low and vice versa. The biological importance of this phenomenon is uncertain.     

Long-distance seed dispersal by wind: disentangling the effects of species traits,vegetation types,vertical turbulence and wind speed

Long-distance dispersal (LDD) of plant seeds by wind is affected by functional traits of the species, specifically seed terminal velocity and height of seed release above the vegetation cover (HAC), as well as by the meteorological parameters wind speed and vertical turbulence. The relative importance of these parameters is still under debate and the importance of their variability in vegetation types, sites and years has only rarely been quantified. To address these topics, we performed simulation studies for different vegetation types, sites, years and plant species with PAPPUS, a process based trajectory model. We found that LDD (measured in terms of migration rates) was higher in forests compared to open landscapes. Forests also showed greater between-year variability in LDD. Terminal velocity had an effect on LDD in both vegetation types, while the effect of HAC was significant only in the open landscape. We found considerable differences in how vertical turbulence and wind speed affect LDD between species and vegetation types: In the open landscape the strength of the positive relationship between vertical turbulence and LDD generally decreases with terminal velocity, whereas it increases in forests. The strength of the predominantly positive effect of wind speed on LDD increases with terminal velocity in both vegetation types, while in forests we found even negative relationships for species with low terminal velocity. Our results generally suggest that the effects of vertical turbulence and wind speed on LDD by wind diverge for species with different functional traits as well as in different vegetation types.     

The relation between maximal running speed and body mass in terrestrial mammals

The available data on maximal running speeds of mammals are presented, and the relationship between speed and body mass is considered. For all mammals ( n = 106), maximal running speed scales as (body mass)^0–17; however, the largest mammals are not the fastest, and an optimal size with regards to running ability is suggested ( 119 kg). Maximal running speeds are, on the average, somewhat more than twice maximal aerobic speeds.
Within the Artiodactyla, Carnivora or Rodentia, maximal running speed is mass independent, in agreement with theoretical expectations for geometrically similar animals (Thompson, 1917; Hill, 1950). McMahon's (1975 b ) model for elastic similarity is therefore not supported by the available data on maximal running speeds, and there appears to be no necessary correspondence between scaling of limb bone proportions and running ability.     

Mechanical model for theoretical determination of maximum running speed in mammals

A mechanical model for the determination of maximum speed in terrestrial tetrapods, designed for application to extinct species, is proposed. Only external bone measures and average body mass estimations are used as input data, and the hypothesis is made that leg bones are strong enough to endure the stress of running at maximum speed at a certain universal safety factor. The model is applied to a broad sample of living mammalian species to test its predictive power, and it is found to provide very good estimates of maximum running speed.     

Sex chromosome-to-autosome transposition events counter Y-chromosome gene loss in mammals

BackgroundAlthough the mammalian X and Y chromosomes evolved from a single pair of autosomes, they are highly differentiated: the Y chromosome is dramatically smaller than the X and has lost most of its genes. The surviving genes are a specialized set with extraordinary evolutionary longevity. Most mammalian lineages have experienced delayed, or relatively recent, loss of at least one conserved Y-linked gene. An extreme example of this phenomenon is in the Japanese spiny rat, where the Y chromosome has disappeared altogether. In this species, many Y-linked genes were rescued by transposition to new genomic locations, but until our work presented here, this has been considered an isolated case.ResultsWe describe eight cases of genes that have relocated to autosomes in mammalian lineages where the corresponding Y-linked gene has been lost. These gene transpositions originated from either the X or Y chromosomes, and are observed in diverse mammalian lineages: occurring at least once in marsupials, apes, and cattle, and at least twice in rodents and marmoset. For two genes - EIF1AX/Y and RPS4X/Y - transposition to autosomes occurred independently in three distinct lineages.ConclusionsRescue of Y-linked gene loss through transposition to autosomes has previously been reported for a single isolated rodent species. However, our findings indicate that this compensatory mechanism is widespread among mammalian species. Thus, Y-linked gene loss emerges as an additional driver of gene transposition from the sex chromosomes, a phenomenon thought to be driven primarily by meiotic sex chromosome inactivation.

Electronic supplementary material

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