Mechanics of running of the ostrich (Struthio camelus)

Ostriches have been filmed running fast in their natural habitat. A female ostrich has been dissected and the principal bones, muscles and tendons in a leg have been measured. It is calculated that stresses up to 240 kN m⁻² and 40 MN m⁻², respectively, act in the digital flexor muscles and their tendons during running. Tensile and compressive stresses up to about 70MNm⁻² and 110 MNm⁻² act in the tibiotarsus. A large proportion of the energy which would otherwise be required for running is probably saved by elastic storage in tendons. Comparisons are made with the legs of flying birds and of antelopes.     

Carbonic anhydrase inhibitors. Inhibition of red blood cell ostrich (Struthio camelus) carbonic anhydrase with a series of aromatic and heterocyclic sulfonamides

The purification of red blood cell carbonic anhydrase (CA, EC 4.2.1.1) from ostrich (scCA) blood is reported, as well as an inhibition study of this enzyme with a series of aromatic and heterocylic sulfonamides. The ostrich enzyme showed a high activity, comparable to that of the human isozyme II, with k_cat of 1.2·10⁶ s^{? 1} and k_cat/K_M of 1.8·10⁷ M^{? 1} s^{? 1}, and an inhibition profile quite different from that of the human red blood cell cytosolic isozymes hCA I and II. scCA has generally a lower affinity for sulfonamide inhibitors as compared to hCA I and II. The only sulfonamide which behaved as a very potent inhibitor of this enzyme was ethoxzolamide (K_I = 3.9 nM) whereas acetazolamide and sulfanilamide behaved as weaker inhibitors (inhibition constants in the range 303–570 nM). Several other aromatic and heterocyclic sulfonamides, mostly derivatives of sulfanilamide, homosulfanilamide, 4-aminoethylbenzenesulfonamide or 5-amino-1,3,4-thiadiazole-2-sulfonamide, showed good affinities for the ostrich enzyme, with K_I values in the range 25–72 nM.     

Tibiofibular junction of the South African ostrich (Struthio camelus australis)

The tibiofibular junction of the South African ostrich (Struthio camelus australis) consists of the Ligg. tibiofibularia caudale, craniale proximale and distale, obliquum, and interosseum. The motion range of the fibula, respective to the tibiotarsus, averaged 35°. This junction is, however, not freely mobile as there is a resting position from which the fibula can be pronated an average of 14° (counterclockwise motion of the right fibula from the proximal view) and supinated an average of 21°. Flexion and inward rotation of the tibiotarsus causes fibular supination. This behavior is induced by the lateral meniscus (which follows the movements of the femur on the plateau of the tibiotarsus) and the femorofibular joint surfaces. As the fibular attachment of the popliteus muscle cannot migrate medially, due to its relative fixation by the femorofibular joint surfaces, fibula supination (tibiotarsus pronation), caused by contraction of the popliteus muscle leads to an inward rotation of the tibiotarsus relative to the femur. The ligaments impede excessive pro- and supination. Except for the Lig. tibiofibulare craniale distale, all ligaments also comprise fibers taut in intermediate positions—the Ligg. obliquum and interosseum each have a fiber bundle that is taut in all positions. Tibiotarsus and fibula have no joint surfaces for a common articulation. Hence, the proximal junction should not be termed “articulation” (especially with regard to the distal “syndesmosis”). © 1996 Wiley-Liss, Inc.     

Differentiation of the acrosomal complex in ostrich (Struthio camelus) spermatids

The acrosomal complex of ostrich sperm consists of a small, cone-shaped acrosome and a slender, cylindrical perforatorium housed within a deep endonuclear canal. The perforatorium is almost exclusively endonuclear in location and is only covered by the acrosome at its point of origin in the apical subacrosomal space. The development of the acrosome is generally similar to that described in other non-passerine birds. Small proacrosomal granules (vesicles) emanating from the Golgi apparatus coalesce to form a large, membrane-bound acrosomal vesicle filled with homogeneous, electron-dense material. The acrosomal vesicle attaches to the nucleus via a shallow depression and subsequently collapses to form the typical cap-like acrosome of non-passerine birds. In ostrich spermatids the endonuclear canal becomes obvious when the collapsed acrosomal vesicle has assumed a dumbbell-shaped appearance. The perforatorium, which originates from moderately electron-dense material contained within the apical subacrosomal space, expands within the deepening endonuclear canal. The material of the perforatorium does not originate in the form of an obvious granule as in chicken and budgerigar spermatids. Indications are that in ostrich spermatids the developing acrosome plays a role in the shaping of the tip of the nucleus. The perforatorium, however, appears to represent a residual structure that has no specifically identified function. © 1996 Wiley-Liss, Inc.     

Hematologic and blood chemistry values of the Masai ostrich (Struthio camelus)

Normal mean values for hematocrit, hemoglobin concentration, erythrocyte and leukocyte counts, hematimetric indices, erythrocyte dimensions, glucose, urea, uric acid, cholesterol, creatinine, total bilirubin, serum aspartate aminotransferase, serum alanine aminotransferase, alkaline phosphatase, creatinine phosphokinase, lactic dehydrogenase, inorganic phosphorus, chloride, total plasma protein, sodium, potassium, calcium, and magnesium were obtained from the blood or plasma of four Masai ostriches (Struthio camelus) when juveniles at 5 mo of age and as adults 1 yr later in the Barcelona Zoo (Spain). Young ostriches had significantly lower concentrations of hematocrit, hemoglobin concentration, calcium, and magnesium, and higher levels of total protein and potassium, than the adult individuals. The rest of the parameters were not significantly different between the two age groups. The data obtained provide reference values for Masai ostriches.     

The isolation and characterization of serum albumin from the ostrich (Struthio camelus)

Ostrich serum albumin (OsSA) was purified by a combination of heat fractionation and polyethylene glycol precipitation. Equilibrium centrifugation revealed a relative molecular mass of 71,666 for the purified monomer, whereas the presence of a dimeric form was confirmed by means of PAGE and SDS-PAGE analysis. Compared to other species, relatively high levels of proline, glycine, isoleucine and histidine together with lowered amounts of half cystine, phenylalanine and arginine were observed in OsSA. A single N-terminal aspartic acid was identified. Isolated chicken adipocytes revealed a significantly lower in vitro lipolytic responsiveness towards added glucagon when OsSA replaced bovine serum albumin (BSA) in the medium (Km = 6.359 and 1.135 nM, Vm = 36.70 and 46.72 nmol/hr/micrograms adipocyte DNA for OsSA and BSA respectively).     

A search for genetic markers associated with egg production in the ostrich (Struthio camelus)

The aim of the current study was to search for genetic markers, microsatellite loci associated with laying performance in ostriches. The material consisted of two groups of ostrich hens characterized by high or low laying performance (over 75 and less than 25 eggs per season, respectively). The investigation covered 30 microsatellite loci characteristic for the ostrich (the CAU group) and led to identification of significant differences in allele and genotype frequencies between the two groups of hens considered. Out of a total of 30 microsatellite loci examined, 28 showed different alleles in relation to analyzed performance groups. In hens of high laying performance (HP group, n = 12), specific alleles occurred in 23 microsatellite loci (40 alleles of 243 identified), while in those of low egg production (LP group, n = 12), they occurred in 22 (51 alleles of 243 identified). The results indicate the usefulness of the microsatellite loci as the potential genetic markers associated with laying performance that can be applied for genetic improvement of ostrich flocks.     

Gas exchange and energy metabolism of the ostrich (Struthio camelus) embryo.

We measured oxygen consumption (V(O(2))) and carbon dioxide emission (V(CO(2))) rates, air-cell gas partial pressures of oxygen (P(A)O(2)) and CO(2) (P(A)CO(2)), eggshell water vapour conductance and energy content of the ostrich (Struthio camelus) egg, 'true hatchling' and residual yolk, and calculated RQ and total oxygen consumption (V(O(2)tot)) for ostrich eggs incubated at 36.5 degrees C and 25% relative humidity. The V(O(2)) pattern showed a drop of approximately 5% before internal pipping. V(O(2)) just prior to internal pipping agrees with allometric calculations. Despite the higher incubation temperature compared to other studies, and the resultant shorter incubation duration (42 days), V(O(2)tot) (91.7 l kg(-1)) was similar to a previously reported value. RQ values during the second half of incubation (approx. 0.68) were lower than expected for lipid catabolism. Prior to internal pipping, P(A)O(2) and P(A)CO(2) were 98 and 48.3 torr (13.1 and 6.4 kPa), respectively. The growth pattern of the ostrich embryo is different from the typical precocial pattern, showing a time delay in the rapid growth phase. As a result, the lowered overall energy expenditure for tissue maintenance, as compared to other species, is reflected in the low yolk utilization and high residual yolk fraction of the whole hatchling dry mass. These could also result from the relatively short incubation period of the ostrich egg, thereby evading desiccation by excess water loss.     

Phylogeographic analysis of nuclear and mtDNA supports subspecies designations in the ostrich (Struthio camelus)

We investigated the phylogeography and subspecies classification of the ostrich (Struthio camelus) by assessing patterns of variation in mitochondrial DNA control region (mtDNA-CR) sequence and across fourteen nuclear microsatellite loci. The current consensus taxonomy of S. camelus names five subspecies based on morphology, geographic range, mtDNA restriction fragment length polymorphism and mtDNA-CR sequence analysis: S. c. camelus, S. c. syriacus, S. c. molybdephanes, S. c. massaicus and S. c. australis. We expanded a previous mtDNA dataset from 18 individual mtDNA-CR sequences to 123 sequences, including sequences from all five subspecies. Importantly, these additional sequences included 43 novel sequences of the red-necked ostrich, S. c. camelus, obtained from birds from Niger. Phylogeographic reconstruction of these sequences matches previous results, with three well-supported clades containing S. c. camelus/syriacus, S. c. molybdophanes, and S. c. massaicus/australis, respectively. The 14 microsatellite loci assessed for 119 individuals of four subspecies (all but S. c. syriacus) showed considerable variation, with an average of 13.4 (±2.0) alleles per locus and a mean observed heterozygosity of 55.7 (±5.3)%. These data revealed high levels of variation within most subspecies, and a structure analysis revealed strong separation between each of the four subspecies. The level of divergence across both marker types suggests the consideration of separate species status for S. c. molybdophanes, and perhaps also for S. c. camelus/syriacus. Both the mtDNA-CR and microsatellite analyzes also suggest that there has been no recent hybridization between the subspecies. These findings are of importance for management of the highly endangered red-necked subspecies (S. c. camelus) and may warrant its placement onto the IUCN red list of threatened animals.     

Isolation and characterization of 70 novel microsatellite markers from ostrich (Struthio camelus) genome.

Microsatellite markers are widely used in linkage mapping, parentage testing, population genetic studies, and molecular evolution studies in many agricultural species, while only a limited number of ostrich (Struthio camelus) microsatellites have been isolated. Thus, we constructed a random small-insert genomic library and a microsatellite-enriched library containing CA repeats. Fourteen clones containing CA repeats were isolated from 3462 clones in the non-enriched library by radioactive screening and 248 positive clones were isolated from 300 sequenced clones from the enriched library by PCR screening. After the enrichment procedures, the proportion of clones containing CA repeats was raised to 78.8%, compared with 0.4% in the non-enriched libraries, indicating that the enrichment value approaches 200 fold, which decreased the time and cost of cloning. The number of complete simple CA repeats in these positive clones ranged from 5 to 29. The primers for 94 of these microsatellites were developed and used to detect polymorphisms, of which 61 loci exhibited length polymorphisms in 17 unrelated ostrich individuals. The new polymorphic microsatellite markers we have identified and characterized will contribute to the ostrich genetic map, parentage testing, and comparative genomics between avian species.     

Phylogeny of neurohypophyseal hormones in birds: microidentification of mesotocin and vasotocin in the ostrich (Struthio camelus)

Ostrich (Struthio camelus) neurohypophysial hormones have been isolated from 5 freeze-dried posterior pituitary glands. Purification has involved three steps: a first molecular sieving on Sephadex G-75 for eliminating proteins, a second molecular sieving on Bio-Gel P4 for separating the two active principles and a high pressure reverse-phase liquid chromatography (HPLC) on Nova-Pak C 18 with an 10 mM acetate-acetonitrile gradient for isolating each hormone. The active peptides have been identified by their retention time in HPLC and their amino acid composition. Mesotocin and vasotocin have thus been characterized. Although the phylogeny of Ratites is disputed, in particular their possible common origin with Carinates, which include most of the living birds, species of the first sub-class seem to have the same neurohypophysial hormones as those of the second.     

Carbonic anhydrase inhibitors. Inhibition of red blood cell ostrich (Struthio camelus) carbonic anhydrase with a series of aromatic and heterocyclic sulfonamides

The purification of red blood cell carbonic anhydrase (CA, EC 4.2.1.1) from ostrich (scCA) blood is reported, as well as an inhibition study of this enzyme with a series of aromatic and heterocylic sulfonamides. The ostrich enzyme showed a high activity, comparable to that of the human isozyme II, with kcat, of 1.2 x 10(6) s(-1) and kcat/KM of 1.8 x 10(7) M(-1)s(-1), and an inhibition profile quite different from that of the human red blood cell cytosolic isozymes hCA I and II. scCA has generally a lower affinity for sulfonamide inhibitors as compared to hCA I and II. The only sulfonamide which behaved as a very potent inhibitor of this enzyme was ethoxzolamide (KI = 3.9 nM) whereas acetazolamide and sulfanilamide behaved as weaker inhibitors (inhibition constants in the range 303-570 nM). Several other aromatic and heterocyclic sulfonamides, mostly derivatives of sulfanilamide, homosulfanilamide, 4-aminoethylbenzenesulfonamide or 5-amino-1,3,4-thiadiazole-2-sulfonamide, showed good affinities for the ostrich enzyme, with KI values in the range 25-72 nM.     

A goose-type lysozyme from ostrich (Struthio camelus) egg white: multiple roles of His101 in its enzymatic reaction

A goose-type lysozyme from ostrich egg white (OEL) was produced by Escherichia coli expression system, and the role of His101 of OEL in the enzymatic reaction was investigated by NMR spectroscopy, thermal unfolding, and theoretical modeling of the enzymatic hydrolysis of hexa-N-acetylchitohexaose, (GlcNAc)₆. Although the binding of tri-N-acetylchitotriose, (GlcNAc)₃, to OEL perturbed several backbone resonances in the ¹H–¹⁵N HSQC spectrum, the chemical shift of the backbone resonance of His101 was not significantly affected. However, apparent pK_a values of His101 and Lys102 determined from the pH titration curves of the backbone chemical shifts were markedly shifted by (GlcNAc)₃ binding. Thermal unfolding experiments and modeling study of (GlcNAc)₆ hydrolysis using a His101-mutated OEL (H101A-OEL) revealed that the His101 mutation affected not only sugar residue affinities at subsites ?3 and ?2 but also the rate constant for bond cleavage. His101 appears to play multiple roles in the substrate binding and the catalytic reaction.     

Studies on avian erythrocyte metabolism: IX. Relationship of changing organic phosphate composition to whole blood oxygen affinity during development of the ostrich (Struthio camelus camelus)

(1) 2,3-Diphosphoglyceric acid (2,3-DPG) is present in erythrocytes (RBC) of the 37-day ostrich embryo at a concentration of 3.7 μmole/ml RBC, representing 23.5% of the cell phosphate. (2) Inositol tetraphosphate (inositol-P₄) is absent in red cells of the 37-day embryo, appears in the RBC about 63 days after hatch, and thereafter gradually accumulates to a level of 2.8 μmole/ml RBC in the adult bird, representing 30–35% of the cell phosphate. (3) Inositol pentaphosphate (inositol-P₅) is present in the erythrocytes of the 37-day embryo at a concentration of 0.8 μmole/ml RBC, reaches a peak level of 2.9 μmole/ml RBC by Day 63 after hatch, and thereafter declines to a level of 1.1 μmole/ml RBC in the adult bird. (4) The crossover time where the molar concentrations of inositol-P₅ and inositol-P₄ are equal in the erythrocyte appears to be about 150 days after hatch. (5) The p₅₀ of whole blood from the 37-day ostrich embryo, 5-day ostrich chick, and adult ostrich was 15.4, 31.8, and 24.9 Torr. The p₅₀ of whole blood immediately after hatch correlates best with an abrupt rise in ATP concentration but after 54 days posthatch correlates best with the appearance of increasing concentrations of inositol-P₄ and the decrease in concentrations of ATP and inositol-P₅. (6) The appearance of inositol-P₄ in the cells could be an adaptive mechanism for regulating oxygen supply by switching to a modulator which maintains a higher oxygen affinity than would a predominance of inositol-P₅.     

Retention of solutes and different-sized particles in the digestive tract of the ostrich (Struthio camelus massaicus), and a comparison with mammals and reptiles

Ostriches (Struthio camelus) achieve digesta retention times, digesta particle size reduction and digestibilities equal to similar-sized herbivorous mammals, in contrast to some other avian herbivores. The sequence of digestive processes in their gastrointestinal tract, however, is still unexplored. Using two groups of four ostriches (mean body mass 75.1 ± 17.3 kg) kept on fresh alfalfa, we tested the effect of two intake levels (17 and 42 g dry matter kg(-0.75)d(-1)) on the mean retention time (MRT) of a solute and three different-sized (2, 10, 20 mm) particle markers, mean faecal particle size (MPS), and digestibility. Intake level did not affect MRT, but MPS (0.74 vs. 1.52 mm) and dry matter digestibility (81 vs. 78%). The solute marker (MRT 22-26 h) was excreted faster than the particle markers; there was no difference in the MRT of 10 and 20 mm particles (MRT 28-32 h), but 2mm particles were retained longer (MRT 39-40 h). Because the solute marker was not selectively retained, and wet-sieving of gut contents of slaughtered animals did not indicate smaller particles in the caeca, the long MRT of small particles is interpreted as intermittent excretion from the gizzard, potentially due to entrapment in small grit. The marker excretion pattern also showed intermittent peaks for all markers in five of the animals, which indicates non-continuous outflow from the gizzard. When adding our data to literature data on avian herbivores, a dichotomy is evident, with ostrich and hoatzin (Opisthocomus hoazin) displaying long MRTs, high digestibilities, and gut capacities similar to mammalian herbivores, and other avian herbivores such as grouse, geese or emus with shorter MRTs, lower fibre digestibilities and lower gut capacities. In the available data for all avian herbivores where food intake and MRTs were measured, this dichotomy and food intake level, but not body mass, was related to MRT, adding to the evidence that body mass itself may not be sole major determinant of digestive physiology. The most striking difference between mammalian and avian herbivores from the literature is the fundamentally lower methane production measured in the very few studies in birds including ostriches, which appears to be at the level of reptiles, in spite of general food intake levels of a magnitude as in mammals. Further studies in ostriches and other avian herbivores are required to understand the differences in digestive mechanisms between avian and mammalian herbivores.     

A comparative study on the histological structure of the spleen in the ostrich (Struthio camelus), the kestrel (Falco tinnunculus) and the osprey (Pandion haliaetus)

The spleen structurally and functionally belongs to the hematopoietic organs and is also an important component of the reticuloendothelial system, which is known to play a major role in host defense. The histological structure of the spleen was investigated in the ostrich, a non-flying bird, the kestrel, a raptor, and the osprey, a fish-eating bird of prey (fish eagle). For this purpose, Mallory's modified triple stain, methyl green-pyronin and silver stain were used. Germinal centers were not present in the spleen of the osprey. In the spleen of the kestrel, penicillar arterioles and the surrounding lymphoid tissue were markedly dense. Compared to the other two birds, the red and white pulps were clearly distinguishable in the spleen of the ostrich.     

Yolk lipids and their fatty acids in the wild and captive ostrich (Struthio camelus)

A comparative study has been made of the major lipid fractions and their fatty acid compositions in the yolk of eggs from ostriches under wild and farmed conditions. There were no differences in the lipid contents and proportions of the lipid fractions between the two groups of yolks. In both groups of yolks triacylglycerol and phospholipid were the major fractions. In the eggs from the wild ostriches, all the lipid fractions displayed substantial concentrations of C18 polyunsaturated fatty acids, triacylglycerol being particularly rich in linolenic acid and phospholipid rich in linoleic acid; phospholipid displayed substantial concentrations also of C20 and C22 polyunsaturates. There were considerable differences in the fatty acid compositions between the yolks. Those from the farmed birds displayed lower proportions of C18 polyunsaturates, particularly linolenic acid, throughout the lipid fractions. Compensatory increases were displayed most obviously in the concentrations of oleic acid and palmitoleic acid as well as other acids. The distinctive and extensive changes in fatty acid composition, particularly relating to the polyunsaturates, are discussed with respect to overall dietary requirements and specificities for embryo metabolism and possible effects on reproductive performance.     

Light and scanning electron microscopic study of the structure of the ostrich (Strutio camelus) tongue

The ostrich's tongue is situated in the posterior part of the oropharyngeal cavity and its length is only about a quarter of the beak cavity. The triangular shortened tongue has retained the usual division into the apex, the body and the root. There are no conical papillae between the body and the root of the tongue, and the presence of the flat fold with lateral processes sliding over the tongue root in the posterior part of the lingual body is a unique morphological feature. All lingual mucosa covers non-keratinised stratified epithelium, and the lamina propria of the mucosa is filled with mucous glands whose round or semilunar openings are found on both the dorsal and ventral surface of the tongue. The complex glands found in the lingual body are composed of alveoli and/or tubules. Moreover, simple tubular glands seen in the posterior part of the tongue root are an exception. Numerous observations have shown that the ostrich's tongue is a modified structure, though not a rudimentary one, whose main function is to produce the secretion moisturising the beak cavity surface and the ingested semidry plant food in this savannah species.