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Amino acid sequences of hemoglobin from guinea fowl (Numida meleagri) and California quail (Lophortyx californica) with phylogenetic analysis of major groups of Galliformes.

We determined the complete amino acid sequences of the hemoglobin of two species, guinea fowl and California quail, in Galliformes from intact globin chain and chemical cleavage fragments in order to analyze the molecular evolution of hemoglobin for the classification of Galliformes. Galliformes have two types of hemoglobin components, HbA and HbD, which consist of identical beta chain and different alpha chains. The sequences are similar to globin chains of Galliformes reported previously. These sequences were compared with those of other Galliformes (Phasianidae, Meleagrididae) using duck and goshawk as out-groups. The phylogenetic tree of major groups of Galliformes based on hemoglobin was similar to the tree model produced based on the amino acid sequence of lysozyme c.

Amino Acid Sequence↗

Calibration of galliform molecular clocks using multiple fossils and genetic partitions.

For more than a century, members of the traditional avian order Galliformes (i.e., pheasants, partridges, junglefowl, and relatives) have been among the most intensively studied birds, but still a comprehensive timeframe for their evolutionary history is lacking. Thanks to a number of recent cladistic interpretations for several galliform fossils, candidates now exist that can potentially be used as accurate internal calibrations for molecular clocks. Here, we describe a molecular timescale for Galliformes based on cytochrome b and ND2 using nine mostly internal fossil-based anchorpoints. Beyond application of calibrations spanning the entire evolutionary history of Galliformes, care was taken to investigate the effects of calibration choice, substitution saturation, and rate heterogeneity among lineages on divergence time estimation. Results show broad consistency in time estimation with five out of the nine total calibrations. Our divergence time estimates, based on these anchorpoints, indicate that the early history of Galliformes took place in the Cretaceous, including the origin of the basal-most megapode and perhaps cracid lineages, but that the remaining morphological diversification likely started in the earliest Tertiary. The multi-calibration/multi-genetic partition approach used here highlights the importance of understanding the genetic saturation, variation, and rate constancy spectra for the accurate calculation of divergence times by use of molecular clocks.

Animals↗

Molecular cloning and characterization of novel centromeric repetitive DNA sequences in the blue-breasted quail (Coturnix chinensis, Galliformes).

A new family of centromeric highly repetitive DNA sequences was isolated from EcoRI-digested genomic DNA of the blue-breasted quail (Coturnix chinensis, Galliformes), and characterized by filter hybridization and chromosome in situ hybridization. The repeated elements were divided into two types by nucleotide length and chromosomal distribution; the 578-bp element predominantly localized to microchromosomes and the 1,524-bp element localized to chromosomes 1 and 2. The 578-bp element represented tandem arrays and did not hybridize to genomic DNAs of other Galliformes species, chicken (Gallus gallus), Japanese quail (Coturnix japonica) and guinea fowl (Numida meleagris). On the other hand, the 1,524-bp element was not organized in tandem arrays, and did hybridize to the genomic DNAs of three other Galliformes species, suggesting that the 1,524-bp element is highly conserved in the Galliformes. The 578-bp element was composed of basic 20-bp internal repeats, and the consensus nucleotide sequence of the internal repeats had homologies to the 41-42 bp CNM repeat and the XHOI family repeat of chicken. Our data suggest that the microchromosome-specific highly repetitive sequences of the blue-breasted quail and chicken were derived from a common ancestral sequence, and that they are one of the major and essential components of chromosomal heterochromatin in Galliformes species.

Animals↗

Karyotypic evolution in the Galliformes: an examination of the process of karyotypic evolution by comparison of the molecular cytogenetic findings with the molecular phylogeny.

To define the process of karyotypic evolution in the Galliformes on a molecular basis, we conducted genome-wide comparative chromosome painting for eight species, i.e. silver pheasant (Lophura nycthemera), Lady Amherst's pheasant (Chrysolophus amherstiae), ring-necked pheasant (Phasianus colchicus), turkey (Meleagris gallopavo), Western capercaillie (Tetrao urogallus), Chinese bamboo-partridge (Bambusicola thoracica) and common peafowl (Pavo cristatus) of the Phasianidae, and plain chachalaca (Ortalis vetula) of the Cracidae, with chicken DNA probes of chromosomes 1-9 and Z. Including our previous data from five other species, chicken (Gallus gallus), Japanese quail (Coturnix japonica) and blue-breasted quail (Coturnix chinensis) of the Phasianidae, guinea fowl (Numida meleagris) of the Numididae and California quail (Callipepla californica) of the Odontophoridae, we represented the evolutionary changes of karyotypes in the 13 species of the Galliformes. In addition, we compared the cytogenetic data with the molecular phylogeny of the 13 species constructed with the nucleotide sequences of the mitochondrial cytochrome b gene, and discussed the process of karyotypic evolution in the Galliformes. Comparative chromosome painting confirmed the previous data on chromosome rearrangements obtained by G-banding analysis, and identified several novel chromosome rearrangements. The process of the evolutionary changes of macrochromosomes in the 13 species was in good accordance with the molecular phylogeny, and the ancestral karyotype of the Galliformes is represented.

Animals↗

Insertion events of CR1 retrotransposable elements elucidate the phylogenetic branching order in galliform birds.

Using standard phylogenetic methods, it can be hard to resolve the order in which speciation events took place when new lineages evolved in the distant past and within a short time frame. As an example, phylogenies of galliform birds (including well-known species such as chicken, turkey, and quail) usually show low bootstrap support values at short internal branches, reflecting the rapid diversification of these birds in the Eocene. However, given the key role of chicken and related poultry species in agricultural, evolutionary, general biological and disease studies, it is important to know their internal relationships. Recently, insertion patterns of transposable elements such as long and short interspersed nuclear element markers have proved powerful in revealing branching orders of difficult phylogenies. Here we decipher the order of speciation events in a group of 27 galliform species based on insertion events of chicken repeat 1 (CR1) transposable elements. Forty-four CR1 marker loci were identified from the draft sequence of the chicken genome, and from turkey BAC clone sequence, and the presence or absence of markers across species was investigated via electrophoretic size separation of amplification products and subsequent confirmation by DNA sequencing. Thirty markers proved possible to type with electrophoresis of which 20 were phylogenetically informative. The distribution of these repeat elements supported a single homoplasy-free cladogram, which confirmed that megapodes, cracids, New World quail, and guinea fowl form outgroups to Phasianidae and that quails, pheasants, and partridges are each polyphyletic groups. Importantly, we show that chicken is an outgroup to turkey and quail, an observation which does not have significant support from previous DNA sequence- and DNA-DNA hybridization-based trees and has important implications for evolutionary studies based on sequence or karyotype data from galliforms. We discuss the potential and limitations of using a genome-based retrotransposon approach in resolving problematic phylogenies among birds.

Animals↗

Comparative growth rates and oxygen consumption in young Galliformes.

The high correlation between growth rate and adult body weight has been much more thoroughly documented for altricial birds than for precocial species. This paper gathers data from the literature for precocial Galliformes and also reports new growth data on six galliform species for analysis. The onset of homeothermic ability is investigated in Galliformes over a range of body size. The results confirm that (1) large species' chicks grow at a slower rate than those of smaller species, and (2) larger species' chicks can thermoregulate earlier than smaller species' chicks under cold stress situations. Published embryonic body weights are also analysed to determine when growth rate differences appear in the development of precocial species. No interspecific differences appeared in the relative growth rates of embryos, and therefore species body size does not appear to influence growth rate before hatching.

Aging↗

Immunoglobulins of the non-galliform birds: antibody expression and repertoire in the duck.

Galliform and non-galliform birds express three immunoglobulin isotypes, IgM, IgA and IgY. Beyond this we should not generalize because differences in gene organization may have functional consequences reflected in the immune response. At present, studies on non-galliform birds are largely restricted to ducks. Ducks express an alternatively spliced form of their IgY heavy chain (upsilon) gene, the IgY(DeltaFc), that lacks the Fc region and Fc-associated secondary effector functions. It is not known how common the expression of the IgY(DeltaFc) is among birds, nor the functional consequences. It is also not known whether the unusual organization of the duck IgH locus, also shared with the chicken, having the gene order of mu, alpha and upsilon, with alpha inverted in the locus, is unique to the galloanseriform lineage. Ducks, like chickens, have a single immunoglobulin light chain of the lambda (lambda) type. Evidence suggests that ducks, like chickens, generate their immunoglobulin repertoire through a single functional rearrangement of the variable (V) region, and generate diversity through gene conversion from a pool of pseudogenes. In Southern blots of germline and rearranged bursal DNA, both the heavy and light chain loci of ducks appear to each undergo one major rearrangement event. For both heavy and light chains, the functional V region element and the pseudogenes appear to consist of a single gene family. Further analysis of 26 heavy chain joining (JH) and 27 light chain JL segments shows there is use of a single J segment in ducks, which is diversified presumably through somatic mutations and gene conversion events. Despite this limitation on the rearrangement of immunoglobulin genes, analysis of 26 DH and 122 VL sequences suggests that extensive sequence diversity is generated.

Animals↗

Phylogeny of Tetraoninae and other galliform birds using mitochondrial 12S and ND2 genes.

The avian subfamily Tetraoninae (grouse and ptarmigan) is a Holarctic group in the order Galliformes distinguished by morphological adaptations to cold environments and behavioral traits associated with elaborate courtship. Here we investigate the relationships of 17 tetraonines and 12 other galliform species using mitochondrial 12S and ND2 sequence data. We found support for the recent phylogenetic classification that separates the genus Dendragapus into two genera, Falcipennis and Dendragapus. In addition, we found support for a tetraonine clade in which the first divergence is between Bonasa umbellus and all others, followed by divergence between a Bonasa bonasia/Bonasa sewerzowi clade and the remaining tetraonines. Falcipennis canadensis is sister to a clade with four Tetrao species, and the genus Centrocercus is sister to a Dendragapus obscurus/Tympanuchus clade. Our data indicate a basal position for Cracidae and Megapodiidae among the five recognized galliform families. We also found strong support for the monophyly of Phasianidae, although the relative positions of Numididae and Odontiphoridae remains unresolved. We use a maximum likelihood approach to infer ages of 37mya for divergence of Numididae and Phasianidae and 28mya for the divergence of Tetraoninae and Meleagris gallopavo. These estimates must be viewed as tentative as they depend on tests of rates of molecular evolution and accurate fossil dates.

Animals↗

Shivering thermogenesis in leg and breast muscles of galliform chicks and nestlings of the domestic pigeon.

We studied the ontogeny of shivering thermogenesis in breast and leg muscles of precocial galliforms (domestic fowl, grey partridge, and Japanese quail) and the altricial domestic pigeon using electromyography (EMG) and indirect calorimetry. Galliforms were able to increase heat production by shivering in leg muscles at the youngest age studied (1-2 d). Pectorals contributed to heat production from days 7-10 onward, but in the partridge and especially in the fowl, shivering by the pectorals was weaker than in the quail. In the pigeon, shivering began in pectorals and legs at 2 and 4 d of age, respectively, and pectorals had clearly the predominant role in thermogenesis. Despite the early beginning of electrical signs of shivering, significant thermogenesis did not appear in the pigeon before the age of 6 d. All galliforms shivered in bursts, like pigeons aged 2-4 d. From the age of 6 d onward, continuous shivering became predominant in the pigeon. In pectorals of 2-6-d-old pigeons, shivering did not increase linearly during decreasing ambient temperature, as in other muscles and species, but started abruptly, at full intensity. Furthermore, in 2-4-d-old pigeons, cooling induced movement activity in legs. The median frequency of shivering EMGs varied (1) with maturation of the muscle, (2) with size of the adult bird, and (3) between altricials and precocials.

Animals↗

Growth rate and maturation of skeletal muscles over a size range of galliform birds.

The relationship between growth rate and development of function in leg and pectoral muscles was studied in four species of galliform birds ranging from 125 g to 18 kg and, for comparison, in an altricial species, the European starling (80 g). An index to neonatal maturity (muscle dry content proportion as a fraction of adult value) was higher in leg than in pectoral muscles and lower in larger than in smaller galliforms. The maturity index was substantially lower in starling neonates. After the first week posthatch, however, the maturity index was highest in larger species. Exponential growth rates decreased linearly with increasing maturity in both pectoral and leg muscles, following similar regressions in all species including the starling. At a particular value of the maturity index, the exponential growth rate was higher in pectoral than in leg muscles. The exponential growth rates of muscles of neonatal large galliforms were lower than expected from their low maturity. This may represent the down-regulation shortly after hatching of the high exponential growth rate needed to reach a large hatching mass in a short incubation period. A slower growth rate immediately posthatch may be necessary if the relatively immature neonatal digestive system cannot deliver nutrients or metabolized energy required for more rapid growth. Smaller species may not be faced with the constraint of rapid growth toward the end of the embryonic period.

Aging↗

Cytogenetic repartition of chicken CR1 sequences evidenced by PRINS in Galliformes and some other birds.

Chicken repeat 1 (CR1) belongs to the non-long repeat class of retrotransposons. Nearly 100000 repeats interspersed in the chicken genome are subdivided into at least six distinct subfamilies, each 300 bp long and all sharing substantial sequence similarity. CR1-like elements were found in genomes from invertebrates to mammals, suggesting their importance for genome structure and/or function. Moreover, numerous data support the hypothesis of their implication in regulation of gene expression. So, the chromosomal distribution of these CR1 sequences in vertebrates is of great interest to improve our knowledge about the genome structure, function and evolution. A comparison of the cytogenetic distribution of CR1 sequences was performed by PRINS using consensus chicken primers on the chromosomes of chicken and species of several bird orders: Galliformes, Anseriformes, Passeriformes and Falconiformes. The study revealed that CR1 repeats are spread over nearly all chicken chromosomes with a higher density on the macrochromosomes and in particular with hot spots on subtelomeric regions of chromosome 1, 2, 3q, 4q, 5q. Their distribution on the macrochromosomes forms a kind of banding pattern, which was not systematically matched with R- or G-banding. This banding pattern appears to be conserved on the chromosomes of the Galliformes studied, irrespective of their karyotypes, rearranged or not. CR1 primers also show similar signals on the chromosomes of birds phylogenetically more distant (Anseriformes, Passeriformes and Falconiformes). This fact confirms the importance of these sequences at the large scale of bird evolution and in the chromosomal structure. The location of CR1 sequences, and in particular of the hot spots, mainly within the richest CG areas are in conformity with the data on an epigenetic role of these highly conserved sequences.

Animals↗

A molecular timescale for galliform birds accounting for uncertainty in time estimates and heterogeneity of rates of DNA substitutions across lineages and sites.

A recent molecular timescale for major lineages of the Galliformes indicated that Megapodiidae and possibly Cracidae, originated in the Cretaceous, while the remaining families originated in the Tertiary. This timescale was based on clock-like evolution in genetic and taxonomic partitions of mitochondrial ND2 and cyt b DNA sequences, and assumed that ordinal diversification of Galloanserae around 90 million years ago and imposed, whenever appropriate, minimum age constraints based on the fossil record. This approach is not ideal, as it did not account for uncertainty in estimating branch lengths and time, including the calibration time, and heterogeneity in the rate of DNA substitution among sites and in different lineages. Furthermore all the information available in the DNA sequences was not included, and may have been affected by stochastic error in individual gene partitions. Here, we present a follow-up analysis by estimating divergence times using a Bayesian framework that accounts for these possible sources of uncertainty. Our results based on combined and separate analyses of mitochondrial DNA sequences comprised of 1756 sites of 12S rDNA, ND2 and cyt b indicated that (1) Megapodiidae and Cracidae, and likely Odontophoridae, originated in the Cretaceous; (2) estimates based on concatenated genes are less affected by stochastic error among sites and less influenced by the phylogenetic signals of individual gene partitions, which are unequally distributed along the phylogenetic tree; and (3) the use of only an external molecular calibration results in lower estimation of most ingroup node ages. We also point out that galliform fossils may not be as useful for point calibrations as was previously suggested, but instead may be better employed as priors for the estimation of node ages under a Bayesian approach.

Animals↗

Do responses of galliform birds vary adaptively with predator size?

Past studies of galliform anti-predator behavior show that they discriminate between aerial and ground predators, producing distinctive, functionally referential vocalizations to each class. Within the category of aerial predators, however, studies using overhead models, video images and observations of natural encounters with birds of prey report little evidence that galliforms discriminate between different raptor species. This pattern suggests that the aerial alarm response may be triggered by general features of objects moving in the air. To test whether these birds are also sensitive to more detailed differences between raptor species, adult chickens with young were presented with variously sized trained raptors (small, intermediate, large) under controlled conditions. In response to the small hawk, there was a decline in anti-predator aggression and in aerial alarm calling as the young grew older and less vulnerable to attack by a hawk of this size. During the same developmental period, responses to the largest hawk, which posed the smallest threat to the young at all stages, did not change; there were intermediate changes at this time in response to the middle-sized hawk. Thus the anti-predator behavior of the adult birds varied in an adaptive fashion, changing as a function of both chick age and risk. We discuss these results in light of current issues concerning the cognitive mechanisms underlying alarm calling behavior in animals.

Adaptation, Psychological↗

An unusual metal-binding cluster found exclusively in the avian breast muscle troponin T of Galliformes and Craciformes.

A repeating metal-binding (Cu2+ > Ni2+ > Zn2+ approximately Co2+) sequence (HE/AEAH)4 has been identified in troponin T isoforms specifically expressed in the breast but not leg muscles of all Galliformes and Craciformes. It is absent in the skeletal and cardiac muscles of mammals and all other avian species investigated. Concentration of the metal-binding sites is adequate to affect free metal levels in the muscle cell and we suggest a possible link between its presence in breast muscle of Galliformes and the high ratio of breast muscle to total body muscle mass and explosive but short-lived flight pattern of these birds. This sequence can be used for a highly selective metal-affinity chromatographic purification of muscle or engineered TnTs even in high salt and/or urea.

Amino Acid Sequence↗

Cospeciation and horizontal transmission of avian sarcoma and leukosis virus gag genes in galliform birds.

In a study of the evolution and distribution of avian retroviruses, we found avian sarcoma and leukosis virus (ASLV) gag genes in 26 species of galliform birds from North America, Central America, eastern Europe, Asia, and Africa. Nineteen of the 26 host species from whom ASLVs were sequenced were not previously known to contain ASLVs. We assessed congruence between ASLV phylogenies based on a total of 110 gag gene sequences and ASLV-host phylogenies based on mitochondrial 12S ribosomal DNA and ND2 sequences to infer coevolutionary history for ASLVs and their hosts. Widespread distribution of ASLVs among diverse, endemic galliform host species suggests an ancient association. Congruent ASLV and host phylogenies for two species of Perdix, two species of Gallus, and Lagopus lagopus and L. mutus also indicate an old association with vertical transmission and cospeciation for these ASLVs and hosts. An inference of horizontal transmission of ASLVs among some members of the Tetraoninae subfamily (grouse and ptarmigan) is supported by ASLV monophyletic groups reflecting geographic distribution and proximity of hosts rather than host species phylogeny. We provide a preliminary phylogenetic taxonomy for the new ASLVs, in which named taxa denote monophyletic groups.

Animals↗

Quantitative development of brain and brain structures in birds (galliformes and passeriformes) compared to that in mammals (insectivores and primates).

The brain weight and brain structure volumes of galliform and passeriform birds were calculated and related to body weight. The total brains and 14 brain regions were investigated in order to calculate factors by which these structures in passeriforms exceed those in galliforms in size. The larger passeriform brains have larger telencephala, especially ventral hyperstriata and neostriata. The enlargement of total brain and telencephalon resembles that in primates, compared to insectivores, within mammals. The enlargement of the ventral hyperstriata + neostriata in passeriforms is fundamentally similar to that of the isocortex in mammals: it reflects an expansion of multimodal integrational capacities, as the ventral hyperstriatum and neostriatum are occupied exclusively or primarily by multimodal integrational areas as is the isocortex.

Animals↗

Haptoglobins in some Galliformes birds.

In several groups of galliform birds (chicken, turkey, pheasant and guinea fowl) the presence and function of plasma haptoglobin (Hp) have been studied. Paper, acetate and starch-gel electrophoresis were carried out and two benzidine stain regions were observed when haemoglobin (Hb) was added to normal plasma (Hb-Hp complex and methemalbumin) besides the free Hb when the binding capacity of these proteins were surpassed. The disappearance of the administered Hb-59Fe was exponential which is evidence of the existence of two different rates. Liver, bone marrow and kidney are the main organs related to the 59Fe uptake. These results on Hp in galliforms and the metabolism of the Hb are concordant with the data in the literature on mammals. This suggests the same common physiological role for Hp.

Animals↗

Viremia in three orders of birds (Anseriformes, Galliformes and Passeriformes) inoculated with Ockelbo virus.

One-hundred six birds of 14 species were inoculated with approximately 10(2.7) plaque-forming units of Ockelbo virus and bled daily for 5 days to determine viremia levels. Virus was detected in birds of all 14 species tested (four Anseriformes, one Galliformes and nine Passeriformes). The onset of viremia occurred earlier and viral titers were higher in very young anseriforms and galliforms than in older birds. Adult passeriforms had Ockelbo viremias of higher titer and longer duration than did adult anseriforms. Viremia titers in adult birds of all three orders tested were sufficient to induce high transmission rates in enzootic mosquito vectors, and viremias in passeriforms could induce high transmission rates in bridging vectors as well. Passeriforms of the genera Turdus and Fringilla could serve as amplification hosts for Ockelbo virus based on the presently demonstrated viremia of high titer and long duration in these birds, and the previously demonstrated high prevalence of Ockelbo virus neutralizing antibodies in free-ranging individuals and great population size compared to birds of other taxa. Bird species of all three orders tested, however, could function as incidental hosts of the virus.

Age Factors↗