The linear alometric relationship between total metabolic energy per life span and body mass of poikilothermic animals

A linear relationship exists between total metabolic energy per life span PT(ls) (kJ) and body mass M (kg) of 54 poikilothermic species (Protozoa, Nematoda, Mollusca, Asteroidae, Insecta, Arachnoidae, Crustacea, Pisces, Amphibia, Reptilia and Snakes): PT(ls) = A(ls*)M(1.0838), where P (kJ/day) is the rate of metabolism and T(ls) (days) is the life span of animals. The linear coefficient A(ls*) = 3.7 x 10(5) kJ/kg is the total metabolic energy, exhausted during the life span per 1 kg body mass of animals. This linear coefficient can be regarded as relatively constant metabolic parameter for poikilothermic organisms, ranging from 0.1 x 10(5) to 5.5 x 10(5) kJ/kg, in spite of 17-degree differences between metabolic rate and body mass of animals. A linear relationship between total metabolic energy per life span and body mass of only 48 poikilothermic multicellular animals (without Protozoa) is: PT(ls) = A(ls*)M(0.9692) with linear coefficient A(ls*) = 2.34 x 10(5) kJ/kg. Since a power relationship exists between the rate of metabolism and body mass of animals of the type: P = aMk (a and k are the alometric constants), an empiric rule could be formulated, that life span is a time interval for which the total metabolic energy per life span becomes proportional to body mass of animals and power coefficient k becomes approximately 1.0.     

Evolution of metatherian and eutherian (mammalian) characters: a review based on cladistic methodology

Cladistic methodology is used to test the hypothesis that three major monophyletic groups exist among living mammals–the oviparous monotremes (Prototheria), and the viviparous marsupials (Metatheria) and placentals (Eutheria). Evaluation is made of the polarity (i.e. the direction of change in a primitive-to-derived sequence) of numerous characters which distinguish some or all of these groups, and of the usefulness of these characters in phylogenetic inference. An attempt is made to establish the state of these characters in the common Late Jurassic-Early Cretaceous therian ancestor of marsupials and placentals.
It is concluded that the most basic division of the Mammalia is the dichotomy into the subclasses Prototheria (including Monotremata, Multituberculata, Triconodonta, Docodonta) and Theria (including Metatheria, Eutheria, Pantotheria and Symmetrodonta). Two major groups exist among living viviparous mammals, the Metatheria and Eutheria; in a cladistic framework these are sister-groups. It is demonstrated that there is no special (sister-group) relationship between monotremes and marsupials, and there is no justification for placing them in a group Marsupionta.     

Relationships Among the Families and Orders of Marsupials and the Major Mammalian Lineages Based on Recombination Activating Gene-1

Controversies remain over the relationships among several of the marsupial families and between the three major extant lineages of mammals: Eutheria (placentals), Metatheria (marsupials), and Prototheria (monotremes). Two opposing hypotheses place the marsupials as either sister to the placental mammals (Theria hypothesis) or sister to the monotremes (Palimpsest or Marsupionta hypothesis). A nuclear gene that has proved useful for analyzing phylogenies of vertebrates is the recombination activation gene-1 (RAG1). RAG1 is a highly conserved gene in vertebrates and likely entered the genome by horizontal transfer early in the evolution of jawed vertebrates. Phylogenetic analyses were performed on RAG1 sequences from seven placentals, 28 marsupials, and all three living monotreme species. Phylogenetic analyses of RAG1 sequences support many of the traditional relationships among the marsupials and suggest a relationship between bandicoots (order Peramelina) and the marsupial mole (order Notoryctemorphia), two lineages whose position in the phylogenetic tree has been enigmatic. A sister relationship between South American shrew opossums (order Paucituberculata) and all other living marsupial orders is also suggested by RAG1. The relationship between the three major groups of mammals is consistent with the Theria hypothesis, with the monotremes as the sister group to a clade containing marsupials and placentals.     

Inference of higher-order relationships in the cycads from a large chloroplast data set

Theria includes Eutheria and its sister taxon Metatheria. Placentalia includes extant eutherians plus their most recent common ancestor. The oldest eutherian is from 125mya (million years ago). Molecular studies place this origin at about 130-185mya. Older dates cannot be refuted based on fossil evidence as earliest eutherian remains are scarce. Earliest superordinal clades (hence Placentalia) range from 64-104mya (median 84mya) based on molecules, similar to 85-90mya based on fossils. Superordinal clades Archonta, Ferungulata, Glires, and Paenungulata based on fossils are similar to molecularly based clades, except Afrotheria was not predicted by fossils. Both fossils and molecules recognize 16 of 18 extant placental orders. Fossils place the origins of orders around 65mya as do some molecular studies, but others suggest ordinal diversification as old as 100mya. Fossil evidence supports a Laurasian origin for Eutheria (and Metatheria) and Placentalia, although some molecular studies suggest a Gondwanan origin for both taxa.     

Marsupials and Eutherians reunited: genetic evidence for the Theria hypothesis of mammalian evolution

The three living monophyletic divisions of Class Mammalia are the Prototheria (monotremes), Metatheria (marsupials), and Eutheria (`placental' mammals). Determining the sister relationships among these three groups is the most fundamental question in mammalian evolution. Phylogenetic comparison of these mammals by either anatomy or mitochondrial DNA has resulted in two conflicting hypotheses, Theria and Marsupionta, and has fueled a ``genes versus morphology' controversy. We have cloned and analyzed a large nuclear gene, the mannose 6-phosphate/insulin-like growth factor II receptor (M6P/IGF2R), from representatives of all three mammalian groups, including platypus, echidna, opossum, wallaby, hedgehog, mouse, rat, rabbit, cow, pig, bat, tree shrew, colugo, ringtail lemur, and human. Statistical analysis of this nuclear gene unambiguously supports the morphology-based Theria hypothesis that excludes monotremes from a clade of marsupials and eutherians. The M6P/IGF2R was also able to resolve the finer structure of the eutherian mammalian family tree. In particular, our analyses support sister group relationships between lagomorphs and rodents, and between the primates and Dermoptera. Statistical support for the grouping of the hedgehog with Feruungulata and Chiroptera was also strong. Received: 8 December 2000 / Accepted: 01 February 2001     

Metabolic costs of growth in free-living Garter Snakes and the energy budgets of ectotherms

1. The metabolic or respiratory cost of growth ( R _G) is the increase in metabolic rate of a growing animal, and it represents chemical potential energy expended in support of net biosynthesis but not deposited as new tissue.
2. Two statistical methods (multiple non-linear regression and analysis of regression residuals) were used to calculate R _G from data ( n = 68) from a doubly labelled water study of free-ranging Garter Snakes ( Thamnophis sirtalis fitchi ) in northern California.
3. The sample-wise ('ecological') cost of growth was 2·07 kJ per gram of net growth (equivalent to 8·63 kJ g^–1 dry tissue); reanalysis of a subset of efficient growers yielded a more conservative 'physiological' estimate of 1·67 kJ g^–1.
4. Our empirical estimate of R _G, among the first reported for squamate reptiles and free-living animals of any kind, compares closely with published, laboratory-derived values for ectotherms.
5. The metabolic costs of growth accounted for an average of 30% of total field metabolic rates for these snakes, which were growing at a mean rate of 3% of body mass per day. However, our method probably underestimated the total ecological cost of growth for large animals, because potential growth costs that covary with body size were not included.
6. Distinction between conceptual and empirical energy budgets clarifies relationships among body size, metabolic rates, and the physiological and ecological costs of growth.     

Comparison of methods for evaluating energy expenditure of incubating wandering albatrosses.

Measurements of incubation energetics can vary depending on the method used to measure metabolism of an incubating bird. Therefore, we evaluated the energy expenditure of six male and four female wandering albatrosses (Diomedea exulans Linnaeus) using doubly labeled water (DLW), the rate of mass loss, and estimates of metabolic water production derived from water influx rate (WIR). Incubation metabolic rates (IMR) determined with DLW (169+/-21 kJ x kg(-1) x d(-1) SD) were significantly lower than estimates derived from mass loss (277+/-46 kJ x kg(-1) x d(-1) SD) and WIR (males=289+/-60 kJ x kg(-1) x d(-1) vs. females=400+/-69 kJ x kg(-1) x d(-1) SD). Estimates of IMR from mass loss and WIR were similar to IMR (305+/-39 kJ x kg(-1) x d(-1) SD) determined by respirometry in a previous study, and IMR from DLW was similar to estimates based on heart rate (HR; 147+/-26 kJ x kg(-1) x d(-1) SD) determined in another study. Applying the different measurements of IMR to construct an energy budget, we estimate that a breeding pair of wandering albatrosses spends 124-234 MJ to incubate the egg for 78 d. Finally, IMRs determined with DLW and HR were similar to estimated basal metabolic rates derived from six different allometric equations, suggesting that heat production from adult maintenance metabolism is sufficient to incubate the egg.     

Loss of genes implicated in gastric function during platypus evolution

Background  

The duck-billed platypus (Ornithorhynchus anatinus) belongs to the mammalian subclass Prototheria, which diverged from the Theria line early in mammalian evolution. The platypus genome sequence provides a unique opportunity to illuminate some aspects of the biology and evolution of these animals.     

Comparative analyses of basal rate of metabolism in mammals: data selection does matter

Basal rate of metabolism (BMR) is a physiological parameter that should be measured under strictly defined experimental conditions. In comparative analyses among mammals BMR is widely used as an index of the intensity of the metabolic machinery or as a proxy for energy expenditure. Many databases with BMR values for mammals are available, but the criteria used to select metabolic data as BMR estimates have often varied and the potential effect of this variability has rarely been questioned. We provide a new, expanded BMR database reflecting compliance with standard criteria (resting, postabsorptive state; thermal neutrality; adult, non‐reproductive status for females) and examine potential effects of differential selectivity on the results of comparative analyses. The database includes 1739 different entries for 817 species of mammals, compiled from the original sources. It provides information permitting assessment of the validity of each estimate and presents the value closest to a proper BMR for each entry. Using different selection criteria, several alternative data sets were extracted and used in comparative analyses of (i) the scaling of BMR to body mass and (ii) the relationship between brain mass and BMR. It was expected that results would be especially dependent on selection criteria with small sample sizes and with relatively weak relationships. Phylogenetically informed regression (phylogenetic generalized least squares, PGLS) was applied to the alternative data sets for several different clades (Mammalia, Eutheria, Metatheria, or individual orders). For Mammalia, a ‘subsampling procedure’ was also applied, in which random subsamples of different sample sizes were taken from each original data set and successively analysed. In each case, two data sets with identical sample size and species, but comprising BMR data with different degrees of reliability, were compared. Selection criteria had minor effects on scaling equations computed for large clades (Mammalia, Eutheria, Metatheria), although less‐reliable estimates of BMR were generally about 12–20% larger than more‐reliable ones. Larger effects were found with more‐limited clades, such as sciuromorph rodents. For the relationship between BMR and brain mass the results of comparative analyses were found to depend strongly on the data set used, especially with more‐limited, order‐level clades. In fact, with small sample sizes (e.g. <100) results often appeared erratic. Subsampling revealed that sample size has a non‐linear effect on the probability of a zero slope for a given relationship. Depending on the species included, results could differ dramatically, especially with small sample sizes. Overall, our findings indicate a need for due diligence when selecting BMR estimates and caution regarding results (even if seemingly significant) with small sample sizes.     

Higher Taxonomic Relationships among Extant Mammals Based on Morphology, with Selected Comparisons of Results from Molecular Data

Until a few decades ago, phylogenetic relationships among placental orders were ambiguous and usually depicted to radiate as an unresolved “bush.” Resolution of this bush by various workers has been progressing slowly, but with promising results corroborated by nondental, dental, and molecular characters. In this study we continue to seek resolution. A total of 258 nondental and 2 dental characters was analyzed by PAUP and MacClade on 39 vertebrate taxa (3 reptiles, 1 nonmammalian therapsid, and 35 mammals; 20 of the mammals are extant and 15 are extinct) to study higher taxonomic relationships with emphasis on Placentalia (Eutheria). About two-thirds of the characters are osteological, the rest concern soft tissues, including myological but excluding molecular characters (most are our data, the rest are from the literature). Cladistic analysis included all 39 taxa (fossil taxa help to evaluate polarities of characters) and all characters were given equal weight. Extant Mammalia are divided into Prototheria and Theria, the latter into Marsupialia and Placentalia. Placentalia comprises Xenarthra and Epitheria. Within Epitheria, Lipotyphla and Preptotheria (emended) are sister-taxa. Preptotherian taxa group into: ungulate-related taxa and various nonungulates. The former include Carnivora, Pholidota, Tubulidentata, Artiodactyla, Cetacea, Perissodactyla, Hyracoidea, Proboscidea, and Sirenia. A possible association to embrace Lagomorpha, Rodentia, Macroscelidea, Scandentia, Primates, Chiroptera, and Dermoptera is suggested. Significant differences between our findings and those of recent investigators include the dissociation of Pholidota from Xenarthra and the plesiomorphous position of Lipotyphla within Epitheria. Congruence between morphological and molecular results is closer than previously reported.     

Fasting metabolism in Antarctic fur seal (Arctocephalus gazella) pups.

The metabolism of 52-73-day old Antarctic fur seal pups from Bird Island, South Georgia, was investigated during fasting periods of normal duration while their mothers were at sea foraging. Body mass decreased exponentially with pups losing 3.5-3.8% of body mass per day. Resting metabolic rate also decreased exponentially from 172-197 ml (O2) x min(-1) at the beginning of the fast and scaled to M(b)(0.74) at 2.3 times the level predicted for adult terrestrial mammals of similar size. While there was no significant sex difference in RMR, female pups had significantly higher (F(1,18)=6.614, P<0.019) mass-specific RMR than male pups throughout the fasting period. Fasting FMR was also significantly (t(15)=2.37, P<0.035) greater in females (823 kJ x kg(-1) x d(-1)) than males (686 kJ x kg(-1) x d(-1)). Average protein turnover during the study period was 19.3 g x d(-1) and contributed to 5.4% of total energy expenditure, indicating the adoption of a protein-sparing strategy with a reliance on primarily lipid catabolism for metabolic energy. This is supported by observed decreases in plasma BUN, U/C, glucose and triglyceride concentrations, and an increase in beta-HBA concentration, indicating that Antarctic fur seals pups adopt this strategy within 2-3 days of fasting. Mean RQ also decreased from 0.77 to 0.72 within 3 days of fasting, further supporting a rapid commencement of protein-sparing. However, RQ gradually increased thereafter to 0.77, suggesting a resumption of protein catabolism which was not substantiated by changes in plasma metabolites. Female pups had higher TBL (%) than males for any given mass, which is consistent with previous findings in this and other fur seal species, and suggests sex differences in metabolic fuel use. The observed changes in plasma metabolites and protein turnover, however, do not support this.     

Energetics of kayaking at submaximal and maximal speeds

The energy cost of kayaking per unit distance (C(k), kJ x m(-1)) was assessed in eight middle- to high-class athletes (three males and five females; 45-76 kg body mass; 1.50-1.88 m height; 15-32 years of age) at submaximal and maximal speeds. At submaximal speeds, C(k) was measured by dividing the steady-state oxygen consumption (VO(2), l x s(-1)) by the speed (v, m x s(-1)), assuming an energy equivalent of 20.9 kJ x l O(-1)(2). At maximal speeds, C(k) was calculated from the ratio of the total metabolic energy expenditure (E, kJ) to the distance (d, m). E was assumed to be the sum of three terms, as originally proposed by Wilkie (1980): E = AnS + alphaVO(2max) x t-alphaVO(2max) x tau(1-e(-t x tau(-1))), were alpha is the energy equivalent of O(2) (20.9 kJ x l O(2)(-1)), tau is the time constant with which VO(2max) is attained at the onset of exercise at the muscular level, AnS is the amount of energy derived from anaerobic energy utilization, t is the performance time, and VO(2max) is the net maximal VO(2). Individual VO(2max) was obtained from the VO(2) measured during the last minute of the 1000-m or 2000-m maximal run. The average metabolic power output (E, kW) amounted to 141% and 102% of the individual maximal aerobic power (VO(2max)) from the shortest (250 m) to the longest (2000 m) distance, respectively. The average (SD) power provided by oxidative processes increased with the distance covered [from 0.64 (0.14) kW at 250 m to 1.02 (0.31) kW at 2000 m], whereas that provided by anaerobic sources showed the opposite trend. The net C(k) was a continuous power function of the speed over the entire range of velocities from 2.88 to 4.45 m x s(-1): C(k) = 0.02 x v(2.26) (r = 0.937, n = 32).     

Body mass and the energy efficiency of locomotion: lessons from incline running.

Rates of oxygen consumption were measured for two bipedal runners (two species of quail) and two quadrupedal runners (two small species of rodents), with average body masses that ranged from 0.035 to 0.217 kg, trained to run on a treadmill set to horizontal and then to a 10 degrees incline. Rates of oxygen consumption increased linearly with speed for all four species and the rates of increase were significantly higher (P < or = 0.05) for all four species when the animals were run on an incline than when they were run on a horizontal. The estimated metabolic energy cost to lift 1 kg mass 1 m vertically was similar for bipeds and quadrupeds of similar body mass and inversely related to body mass for both running styles. When the data for the animals used in the present study are combined with similar data for adult animals from the literature, the results show that the metabolic energy efficiencies of locomotion, estimated from the cost of vertical work, are the same for bipedal and quadrupedal runners. In both groups, the metabolic energy efficiency of locomotion is directly related to body mass for animals smaller than 1 kg body mass.     

Body mass and the energy efficiency of locomotion: lessons from incline running

Rates of oxygen consumption were measured for two bipedal runners (two species of quail) and two quadrupedal runners (two small species of rodents), with average body masses that ranged from 0.035 to 0.217 kg, trained to run on a treadmill set to horizontal and then to a 10 degrees incline. Rates of oxygen consumption increased linearly with speed for all four species and the rates of increase were significantly higher (P < or = 0.05) for all four species when the animals were run on an incline than when they were run on a horizontal. The estimated metabolic energy cost to lift 1 kg mass 1 m vertically was similar for bipeds and quadrupeds of similar body mass and inversely related to body mass for both running styles. When the data for the animals used in the present study are combined with similar data for adult animals from the literature, the results show that the metabolic energy efficiencies of locomotion, estimated from the cost of vertical work, are the same for bipedal and quadrupedal runners. In both groups, the metabolic energy efficiency of locomotion is directly related to body mass for animals smaller than 1 kg body mass.     

Permanent electric polarization and pyroelectric behaviour of the vertebrate skeleton

Summary The segmental organisation of the axial skeleton of the Vertebrata according to sclerotomes and scleromites is clearly evident in the Mammalia by their varying directions of electric polarization. In the corpora vertebrae this is demonstrable until the end of the growth period, in the disci intervertebrales the phenomenon persists through life, and in the ligamenta longitudinalia communia columnae vertebralis it probably persists through life. Two types of vertebral epiphyses were found in the Mammalia.Type I has one single, continuous direction of electric polarization like, for instance, the cartilaginous plates of the vertebrae. Examples of this type are the vertebral epiphyses of the Prototheria (Monotremata) and the Metatheria (Marsupialia), the epiphyses hi the caudal vertebrae of numerous Eutheria (Placentalia) and also the bony vertebral epiphyses of man. In vertebrae having this type of epiphysis, the direction of electric polarization is reversed by 180° at the end of the growth period, both in the epiphysis itself and in the bony centre of the corpus to which it belongs.Type II consists of two layers having opposite directions of electric polarization and is comparable to the epiphyses of certain long bones. This type is found in the trunk vertebrae of the Eutheria, but it was also detected in the caudal vertebrae of the Edentata and Cetacea specimens investigated by us. (Type II occurs neither in man nor probably in the anthropoid apes).In the bony parts of the axial skeleton of the Mammalia, the electric polarization phenomena fade with age; this fading process sets in sooner or later after termination of the growth period. It invariably begins at the centre of the vertebral body and progresses, simultaneously in the two vertebral halves, towards the cranial and caudal end faces of the vertebrae. In old specimens, some weak electric polarization is normally only left in the cranial and caudal cortex. Polarization does not fade with age in the purely collagenous structures of the axial skeleton (e. g. the disci intervertebrales and the ligamenta longitudinalia communia); the polarization phenomena merely become a little less intense at a very great age, but the reduction in intensity varies greatly from one individual to another, and is probably dependent on their vitality.We investigated fossile bone material with the task of determining whether the electric polarization phenomena in the collagenous structures are maintained over long periods of time. Cervical vertebrae, thoracic vertebrae and lumbar vertebrae of juvenile and adult specimens of the cave bear (Ursus spelaeus), which lived in the glacial period known as the Würm-Eiszeit, exhibited electric polarization features that were analogous to those found in contemporary Mammalia of comparable age.The phylogenetic relationship between the Mammalia and certain Reptilia is clearly brought out by the similarity of electric polarization phenomena in the axial skeleton of the Prototheria (Monotremata) and the Crocodilia.

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Zusammenfassung Durch unterschiedliche Richtungen ihrer elektrischen Polarisation tritt bei den Mammalia die segmentale Gliederung des Vertebrata-Achsenskeletts nach Sclerotomen und Scleromiten deutlich hervor. Dies ist der Fall in den Corpora vertebrae bis zum Wachstumsabschluß, in den Processus vertebrae im juvenilen Stadium, in den Disci intervertebrales zeitlebens und in den Ligamenta longitudinalia communia columnae vertebralis wahrscheinlich zeitlebens. Es wurden zwei Typen von Wirbelepiphysen bei den Mammalia festgestellt.Typ I hat nur eine einzige, durchgehende elektrische Polarisationsrichtung, ähnlich wie z. B. die Knorpelplatten der Wirbel. Diesem Typ gehören die Wirbelepiphysen der Prototheria (Monotremata) und der Metatheria (Marsupialia), sowie die Schwanzwirbelepiphysen zahlreicher Eutheria (Placentalia) an, übrigens auch die knöchernen Wirbelepiphysen des Menschen. Bei Wirbeln mit diesem Epiphysentyp findet beim Wachstumsabschluß eine Umkehrung der elektrischen Polarisationsrichtungen um 180° sowohl in den Epiphysen selbst als auch im zugehörigen Wirbelkörperkern statt.Typ II besteht aus zwei Schichten mit einander entgegengesetzten elektrischen Polarisationsrichtungen und hat Ähnlichkeit mit den Epiphysen mancher Röhrenknochen. Er findet sich in den Rumpfwirbeln der Eutheria, wurde aber auch in den Schwanzwirbeln der untersuchten Edentata und Cetacea festgestellt. (Beim Menschen und wahrscheinlich bei den anthropoiden Affen kommt Typ II nicht vor.)Es tritt ein Altersschwund der elektrischen Polarisationserscheinungen in den knöchernen Teilen des Achsenskeletts der Mammalia früher oder später nach dem Wachstumsabschluß auf. Der Altersschwund beginnt stets in der Corpusmitte und setzt sich, in beiden Wirbelhälften gleichzeitig, nach den cranialen und caudalen Wirbelendflächen zu fort. Alte Individuen zeigen meist nur noch in der cranialen und caudalen Cortex eine schwache elektrische Polarisation. Die reinen Kollagenstrukturen des Achsenskeletts (u.a. die Disci intervertebrales und die Ligamenta longitudinalia communia) zeigen keinen Altersschwund, sondern nur eine geringe Abschwächung der elektrischen Polarisationserscheinungen im höchsten Lebensalter und mit ausgeprägter individueller, wahrscheinlich vitalitätsbedingter Unterschiedlichkeit.Um zu prüfen, ob die elektrischen Polarisationserscheinungen von Kollagenstrukturen sich auch im Laufe langer Zeiträume erhalten, wurde fossiles Knochenmaterial untersucht. Hals-, Brust- und Lendenwirbel juveniler und adulter Exemplare des Höhlenbären (Ursus spelaeus) aus der Wurm-Eiszeit zeigten analoge elektrische Polarisationsverhältnisse wie Mammalia vergleichbaren Alters von heute.Die stammesgeschichtliche Verwandtschaft der Mammalia mit bestimmten Reptilia kommt in einer Gleichartigkeit der elektrischen Polarisationsverhältnisse im Achsenskelett der Prototheria (Monotremata) und der Crocodilia zum Ausdruck.

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For Prof. Dr. W. Bargmann.

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With support from the Deutsche Forschungsgemeinschaft which is gratefully acknowledged.     

Effect of undernutrition in foetal life on energy expenditure during gestation in ewes

The long-term effect of early life undernutrition on late gestation energy expenditure (EEgest) was investigated in sheep. Ewes were fed either adequate (100%) or restricted (60%) energy and protein during late foetal life as well as during last trimester of gestation later in life, resulting in three groups: Adequate-Adequate (AA, n = 5), Adequate-Restricted (AR, n = 5) and Restricted-Restricted (RR, n = 5). At two weeks pre-partum, EEgest were calculated from respiratory gaseous exchange and nitrogen excreted in urine and further it was partitioned to energy expenditure for conceptus development (EEconceptus) and homeorhetic adaptations in maternal metabolism (EEhomeorhetic). Late gestational energy and protein restriction reduced the EEgest in the AR ewes (4.1 MJ x d(-1)) but not in the RR ewes (5.2 MJ x d(-1)) compared with the AA ewes (6.8 MJ x d(-1)). Based on conceptus-weight, no significant difference was found in EEhomeorhetic among the groups; 172, 175 and 169 kJ/kg x d(-1) in AA, AR and RR ewes, respectively. However, EEconceptus was significantly lower in AR (135 kJ/kg x d(-1)) in comparison with AA (298 kJ/kg x d(-1)) and RR (252 kJ/kg x d(-1)) ewes. In conclusion, exposure nutrient restriction in early life impairs the ability of ewes to respond to nutritional restriction in terms of energy expenditure of gestation.     

Mammalian genome evolution: new clues from comparisons of eutherians, marsupials and monotremes.

1. Comparisons of chromosomes and gene maps of different mammals are yielding a big picture of the evolution of mammalian genome form and function. It has been particularly instructive to compare gene arrangements on the sex chromosomes between the three major groups of mammals. Eutheria (so-called placental mammals). Metatheria (marsupials) and Prototheria (monotremes), which diverged 150 and 170 Myr BP respectively. 2. A region amounting to 3% of the haploid genome is located on the X chromosome in all three groups, implying that this region must have been part of the original X in a common ancestor. This region comprises the long arm of the human X. 3. A region represented by the short arm of the human X is common to the X in all eutherians, but is autosomal in marsupials and monotremes; thus it was not a part of the original X, and must have been acquired by the X early in the eutherian radiation. 4. This recently acquired region was probably translocated to a pseudoautosomal region shared by the eutherian X and Y. Thus it was originally paired and exempt from X chromosome inactivation; stepwise deletion of this region from the Y and recruitment of the newly unpaired region of the X into the inactivation system could account for some of the peculiarities of this region of the human X. 5. The sex-determining gene TDF must lie on the Y in all mammals in which the Y is male determining. The autosomal location of the candidate gene ZFY in marsupials and monotremes eliminates it from consideration. The recently described candidate gene SRY has yet to pass the "marsupial test".     

Brain gangliosides in monotremes, marsupials and placentals: phylogenetic and thermoregulatory aspects

The concentration and composition of brain gangliosides of 17 mammalian species belonging to the subclasses of Prototheria (monotremes), Metatheria (marsupials), and Eutheria (placentals) were investigated. The mean concentration of brain gangliosides ranges from 525 to 610 micrograms NeuAc/g wet wt in monotremes, 445-900 micrograms in marsupials and from 630 to 1130 micrograms in the placentals. In the phylogenetic series of mammals, a decrease in the complexity of brain ganglioside composition becomes obvious: a drastic reduction in the number of individual ganglioside fractions particularly those of the c-pathway of biosynthesis, took place from the level of monotremes to that of the marsupials and placentals. In monotremes, marsupials and "lower" placentals (insectivores) the percentage of alkali-labile gangliosides is relatively low (between traces and 5%), whereas in the higher evolved mammals it amounts to about 20% of all gangliosides. The ratio of the contents of the two major mammalian ganglioside fractions GD1a and GT1b is generally in the range of 1.0 and even higher; in the heterothermic platypus from the monotremes and in hibernators among the placental mammals, however, it is much lower (about 0.8). These data support the hypothesis that the brain ganglioside composition not only depends on the phylogenetic level of nervous organization (cephalization) but is additionally correlated with the state of thermal adaptation.     

Quantitative characteristics of nutrition in free-ranging reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>) and musk oxen (<Emphasis Type="Italic">Ovibos moschatus</Emphasis>) on Wrangel Island

In 2004 to 2006, studies on free-ranging reindeer and musk oxen on Wrangel island were performed to estimate the nutrient value, daily intake, and digestibility of forage in different seasons of the year. Forage digestibility was determined from the ratio between the contents of indigestible components (lignin) in forage and animal feces. Daily forage intake was calculated from data on daily feces excretion and forage digestibility. The amount of feces and parameters of life activity of individual animals were estimated by following their 24-hour tracks. The results show that the daily intake of forage (dry weight) in musk oxen amounts to 7.8 kg in the summer period and decreases to 6.1 kg in the snow period (March–April). Grazing reindeer in the snow period consume 3.9 kg of forage per day. Forage digestibility in reindeer reaches 56% in March to April and decreases to 52% in June. In musk oxen, forage digestibility in different seasons (March, June, September) varies within a similar range (53–57%). Nutrition parameters (diet composition, forage intake rate and digestibility) of reindeer and musk oxen on grazing grounds of Wrangel Island are similar. Metabolizable energy consumption in both animals during the winter period is at the maintenance level (917–930 kJ/kg0.75 body weight). In the summer period, this parameter in musk oxen increases to 1163 kJ/kg0.75 body weight to meet the energy demand of the animals.     

Effects of obesity and sex on the energetic cost and preferred speed of walking.

The metabolic energy cost of walking is determined, to a large degree, by body mass, but it is not clear how body composition and mass distribution influence this cost. We tested the hypothesis that walking would be most expensive for obese women compared with obese men and normal-weight women and men. Furthermore, we hypothesized that for all groups, preferred walking speed would correspond to the speed that minimized the gross energy cost per distance. We measured body composition, maximal oxygen consumption, and preferred walking speed of 39 (19 class II obese, 20 normal weight) women and men. We also measured oxygen consumption and carbon dioxide production while the subjects walked on a level treadmill at six speeds (0.50-1.75 m/s). Both obesity and sex affected the net metabolic rate (W/kg) of walking. Net metabolic rates of obese subjects were only approximately 10% greater (per kg) than for normal-weight subjects, and net metabolic rates for women were approximately 10% greater than for men. The increase in net metabolic rate at faster walking speeds was greatest in obese women compared with the other groups. Preferred walking speed was not different across groups (1.42 m/s) and was near the speed that minimized gross energy cost per distance. Surprisingly, mass distribution (thigh mass/body mass) was not related to net metabolic rate, but body composition (% fat) was (r2= 0.43). Detailed biomechanical studies of walking are needed to investigate whether obese individuals adopt novel energy saving mechanisms during walking.