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Terrestrial animals not only need to walk and run but also lie prone to rest and then stand up. Sit-to-stand (STS) and sit-to-walk (STW) transitions are vital behaviours little studied in species other than humans so far, but likely impose biomechanical constraints on limb design because they involve near-maximal excursions of limb joints that should require large length changes and force production from muscles. By integrating data from experiments into musculoskeletal simulations, we analysed joint motions, ground reaction forces, and muscle dynamics during STS and STW in a large terrestrial, bipedal and cursorial bird: the emu (Dromaius novaehollandiae; body mass ∼30 kg). Simulation results suggest that in both STS and STW, emus operate near the functional limits (∼50% of shortening/lengthening) of some of their hindlimb muscles, particularly in distal muscles with limited capacity for length change and leverage. Both movements involved high muscle activations (>50%) and force generation of the major joint extensor muscles early in the transition. STW required larger net joint moments and non-sagittal motions than STS, entailing greater demands for muscle capacity. Whilst our study involves multiple assumptions, our findings lay the groundwork for future studies to understand, for example, how tendon contributions may reduce excessive muscle demands, especially in the distal hindlimb. As the first investigation into how an avian species stands up, this study provides a foundational framework for future comparative studies investigating organismal morphofunctional specialisations and evolution, offering potential robotics and animal welfare applications., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

In Australia, significant shifts in species distribution have occurred with the loss of megafauna, changes in indigenous Australian fire regime and land-use changes with European settlement. The emu, one of the last megafaunal species in Australia, has likely undergone substantial distribution changes, particularly near the east coast of Australia where urbanisation is extensive and some populations have declined. We modelled emu distribution across the continental mainland and across the Great Dividing Range region (GDR) of eastern Australia, under historical, present and future climates. We predicted shifts in emu distribution using ensemble modelling, hindcasting and forecasting distribution from current emu occurrence data. Emus have expanded their range northward into central Australia over the 6000 years modelled here. Areas west of the GDR have become more suitable since the mid-Holocene, which was unsuitable then due to high precipitation seasonality. However, the east coast of Australia has become climatically sub-optimal and will remain so for at least 50 years. The north east of NSW encompasses the range of the only listed endangered population, which now occurs at the margins of optimal climatic conditions for emus. Being at the fringe of suitable climatic conditions may put this population at higher risk of further decline from non-climatic anthropogenic disturbances e.g. depredation by introduced foxes and pigs. The limited scientific knowledge about wild emu ecology and biology currently available limits our ability to quantify these risks.

Studies were carried out to determine the influence of age and sex on two types of fat (back fat and abdominal fat) in the emu, as these are factors that influence the composition of animal tissues. The material involved 26 emus at the age 1 (6 males), 3 (6 males) and 15 years (8 females and 6 males), kept on the same farm and fed the feed of the same nutritional value. The basic chemical composition, cholesterol and mineral content, as well as fatty acid profile of back and abdominal fat of emu were determined. Abdominal fat was characterized by higher content of fat and ash, as well as Mn and Ba. Back fat, on the other hand, showed a higher level of protein, cholesterol, C16:1 and the elements K, P, Si, Na, Ca, Mg, Fe, Zn, Se and Cu. With age, regardless of the type of fat tissue, fat content decreased and water content increased. The highest content of protein, ash, cholesterol, some fatty acids (C18:0, C18:1n9c, C18:2n6c), generally higher content of MUFA, PUFA and the elements K, P, Ca, Mg, Fe, Zn, Pb, Se, Cr, Cd, were found in the fatty tissue of 15-year-old emus. Sex did influence the content of Si, Ca, Cu, Sr, which was higher in the fatty tissue of males. The composition of emu storage fat is determined by factors such as age, sex and the location of the fat tissue in the body.

Many ideas have been put forward for the adaptive value of the cassowary casque; and yet, its purpose remains speculative. Homeothermic animals elevate body temperature through metabolic heat production. Heat gain must be offset by heat loss to maintain internal temperatures within a range for optimal performance. Living in a tropical climate, cassowaries, being large bodied, dark feathered birds, are under thermal pressure to offload heat. We tested the original hypothesis that the casque acts as a thermal window. With infrared thermographic analyses of living cassowaries over an expansive range of ambient temperatures, we provide evidence that the casque acts as a thermal radiator, offloading heat at high temperatures and restricting heat loss at low temperatures. Interestingly, at intermediate temperatures, the casque appears thermally heterogeneous, with the posterior of the casque heating up before the front half. These findings might have implications for the function of similar structures in avian and non-avian dinosaurs.

In this study, we analyzed selected morphological traits of eggs, as well as structure, strength, and protein composition of the vitelline membrane (VM) of ostrich, emu, and greater rhea eggs. Ninety eggs (30 for species) were analyzed for the following parameters: egg weight, yolk weight, yolk ratio, and yolk index. In addition, pH value, water activity, consistency index, and flow behavior index were determined. The strength of VM was measured using the TA.HDPlus Texture Analyzer. Micrograph images were taken via a scanning electron microscope. Polyacrylamide gel electrophoresis was conducted under denaturing conditions. Ostrich eggs were characterized by the highest egg and yolk weight compared with those of emu and greater rhea eggs, whereas emu eggs had the highest yolk ratio compared with those of ostrich and greater rhea eggs (P > 0.05). Yolk content differed among the species in terms of water activity; it was found to be higher in emu eggs than in ostrich and greater rhea eggs (P > 0.05). Based on flow curves, yolks of the ratites were classified as pseudoplastic non-Newtonian fluids. The consistency index was significantly higher in yolks of ostrich and emu than that of greater rhea eggs, whereas the VM of yolks of greater rhea eggs was the most resistant (had the highest breaking force = 26.4 g). All species differed significantly regarding the structure of VM, the outer layer (OL) in particular, which was found to constitute fibers of various thicknesses that were differently arranged. Fibers of the OL of the VM of emu, whose fibers were the least differentiated but formed the most compact network, were the most diverse in characterization. An electropherogram of the VM of ostrich revealed 11 primary protein bands: 6 for the OL and 5 for the inner layer (IL), that of emu revealed 9 bands: 5 for the OL and 4 for the IL, and that of greater rhea revealed 10 bands: 6 for the OL and 4 for the IL.

1. The aim of this study was to present selected elements of emu behaviour in three different seasons (the non-breeding season, the pre-breeding season and the breeding season) and in the morning and afternoon. The study focused on feed, water and sand intake; plumage care; resting; agonistic behaviour and other activities, mostly standing still, aimless walking or observing the surroundings. 2. The troop consisted of 11 emus (5 females and 6 males) aged 12 years, kept in a free-range system. Observations were carried out during a period of 3 weeks in each season, during which 60-min observation sessions took place on 3 randomly chosen days. 3. A significant effect was found of each season on the behaviour patterns related to feeding, sand intake, plumage care, resting and agonistic behaviour. 4. Time of day affected the behaviour associated with the collection of food and water, foraging, pecking and ingesting sand and plumage preening. 5. Most of the time, regardless of season and time of day, the emus were standing still, walking aimlessly or observing the surrounding area.

Understanding of the diversity of skeletal loading regimes in vertebrate long bones during locomotion has been significantly enhanced by the application of planar strain theory (PST) to in vivo bone strain data. PST is used to model the distribution of longitudinal strains normal to the bone's transverse cross-section and the location of the neutral axis of bending. To our knowledge, the application of this theory to skeletal biomechanics has not been experimentally validated. We evaluated the accuracy of PST using strain measurements from emu tibiotarsi instrumented with four strain gauges and loaded in ex vivo four-point bending. Using measured strains from three-gauge combinations, PST was applied to predict strain values at a fourth gauge's location. Experimentally measured and predicted strain values correlated linearly with a slope near 1.0, suggesting that PST accurately predicts longitudinal strains. Additionally, we assessed the use of PST to extrapolate shear strains to locations on a bone not instrumented with rosette strain gauges. Guineafowl tibiotarsi were instrumented with rosette strain gauges and in vivo longitudinal and shear strains were measured during treadmill running. Individual-specific and sample-mean ratios between measured longitudinal strains from the medial and posterior bone surfaces were used to extrapolate posterior-site shear strain from shear strains measured on the medial surface. Measured and predicted shear strains at the posterior gauge site using either ratio showed trends for a positive correlation between measured and predicted strains, but the correlation did not equal 1.0 and had a non-zero intercept, suggesting that the use of PST should be carefully considered in the context of the goals of the study and the desired precision for the predicted shear strains., (© 2016. Published by The Company of Biologists Ltd.)

A comprehensive description of the spectral characteristics of retinal photoreceptors in palaeognaths is lacking. Moreover, controversy exists with respect to the spectral sensitivity of the short-wavelength-sensitive-1 (SWS1) opsin-based visual pigment expressed in one type of single cone: previous microspectrophotometric (MSP) measurements in the ostrich (Struthio camelus) suggested a violet-sensitive (VS) SWS1 pigment, but all palaeognath SWS1 opsin sequences obtained to date (including the ostrich) imply that the visual pigment is ultraviolet-sensitive (UVS). In this study, MSP was used to measure the spectral properties of visual pigments and oil droplets in the retinal photoreceptors of the emu (Dromaius novaehollandiae). Results show that the emu resembles most other bird species in possessing four spectrally distinct single cones, as well as double cones and rods. Four cone and a single rod opsin are expressed, each an orthologue of a previously identified pigment. The SWS1 pigment is clearly UVS (wavelength of maximum absorbance [λmax] = 376 nm), with key tuning sites (Phe86 and Cys90) consistent with other vertebrate UVS SWS1 pigments. Palaeognaths would appear, therefore, to have UVS SWS1 pigments. As they are considered to be basal in avian evolution, this suggests that UVS is the most likely ancestral state for birds. The functional significance of a dedicated UVS cone type in the emu is discussed., (© 2016 The Author(s).)

The sutures of the skulls of vertebrates are generally open early in life and slowly close as maturity is attained. The assumption that all vertebrates follow this pattern of progressive sutural closure has been used to assess maturity in the fossil remains of non-avian dinosaurs. Here, we test this assumption in two members of the Extant Phylogenetic Bracket of the Dinosauria, the emu, Dromaius novaehollandiae and the American alligator, Alligator mississippiensis, by investigating the sequence and timing of sutural fusion in their skulls. As expected, almost all the sutures in the emu skull progressively close (i.e., they get narrower) and then obliterate during ontogeny. However, in the American alligator, only two sutures out of 36 obliterate completely and they do so during embryonic development. Surprisingly, as maturity progresses, many sutures of alligators become wider in large individuals compared to younger, smaller individuals. Histological and histomorphometric analyses on two sutures and one synchondrosis in an ontogenetic series of American alligator confirmed our morphological observations. This pattern of sutural widening might reflect feeding biomechanics and dietary changes through ontogeny. Our findings show that progressive sutural closure is not always observed in extant archosaurs, and therefore suggest that cranial sutural fusion is an ambiguous proxy for assessing maturity in non-avian dinosaurs.

Species distribution modeling has been widely used in studying habitat relationships and for conservation purposes. However, neglecting ecological knowledge about species, e.g. their seasonal movements, and ignoring the proper environmental factors that can explain key elements for species survival (shelter, food and water) increase model uncertainty. This study exemplifies how these ecological gaps in species distribution modeling can be addressed by modeling the distribution of the emu (Dromaius novaehollandiae) in Australia. Emus cover a large area during the austral winter. However, their habitat shrinks during the summer months. We show evidence of emu summer habitat shrinkage due to higher fire frequency, and low water and food availability in northern regions. Our findings indicate that emus prefer areas with higher vegetation productivity and low fire recurrence, while their distribution is linked to an optimal intermediate (~0.12 m3 m(-3)) soil moisture range. We propose that the application of three geospatial data products derived from satellite remote sensing, namely fire frequency, ecosystem productivity, and soil water content, provides an effective representation of emu general habitat requirements, and substantially improves species distribution modeling and representation of the species' ecological habitat niche across Australia.

In developing avian embryos, the right and left ductus arteriosi (DA) allow for a shunt of systemic venous return away from the lungs to the body and chorioallantoic membrane (CAM). Unlike in mammals where the transition from placental respiration to lung respiration is instantaneous, in birds the transition from embryonic CAM respiration to lung respiration can take over 24h. To understand the physiological consequences of this long transition we examined circulatory changes and DA morphological changes during hatching in the emu (Dromaius novaehollandiae), a primitive ratite bird. By tracking microspheres injected into a CAM vein, we observed no change in DA blood flow between the pre-pipped to internally pipped stages. Two hours after external pipping, however, a significant decrease in DA blood flow occurred, evident from a decreased systemic blood flow and subsequent increased lung blood flow. Upon hatching, the right-to-left shunt disappeared. These physiological changes in DA blood flow correspond with a large decrease in DA lumen diameter from the pre-pipped stages to Day 1 hatchlings. Upon hatching, the right-to-left shunt disappeared and at the same time apoptosis of smooth muscle cells began remodeling the DA for permanent closure. After the initial smooth muscle contraction, the lumen disappeared as intimal cushioning formed, the internal elastic lamina degenerated, and numerous cells underwent regulated apoptosis. The DA closed rapidly between the initiation of external pipping and hatching, resulting in circulatory patterns similar to the adult. This response is most likely produced by increased DA constriction in response to increased arterial oxygen levels and the initiation of vessel remodeling., (Copyright © 2015 Elsevier Inc. All rights reserved.)

Background: A novel transient structure was observed in the spermatids of three ratite species using transmission electron microscopy., Results: The structure first appeared at the circular manchette stage of sperm development, was most prominent during the longitudinal manchette phase and disappeared abruptly prior to spermiation. It was composed of regularly-spaced finger-like projections which were closely associated with the outer nuclear membrane, giving the nucleus a cogwheel-like appearance. The projections were approximately 30 nm long and 14 nm wide. Although a similar structure has been described in certain lizard and crocodile species, this is the first report of a similar structure in the developing spermatids of birds., Conclusions: The potential value of non-traditional characters, such as spermiogenesis and sperm ultrastructure, as phylogenetic markers has recently been advocated. The morphologically unique structure found in ratite spermatids provides additional evidence of a possible phylogenetic link between the reptiles and birds. It also endorses the basal positioning of the ratites as a monophyletic group within the avian phylogenetic tree.

Blood profiling is a helpful tool in detecting the health status, metabolic diseases, nutritional deficiencies, and welfare of animals. Body weights, body temperatures, hematological and serum biochemical parameters, enzymes, and electrolytes in both sexes of farm emus at the beginning of their breeding season (November in Canada), were determined. The reference interval for each analyte was also calculated. Emus have lower body temperatures (37.2 ± 0.2) than other poultry species. There was no significant between-sex difference in BW, body temperature, and all the hematological and enzyme parameters measured. However, females had significantly (P < 0.001) higher serum calcium, phosphorus, albumin, total protein, globulin, and triglyceride levels than males, probably in preparation for egg laying. We also examined our findings in light of their sex-role reversal in incubation and brooding. Contrary to other avian species in which only females incubate and brood, there was no sex difference in the hematological and enzyme parameters measured in emus. We found that emus are similar to other ratite species with respect to the changes in protein, globulin, triglyceride, and calcium levels. The findings from our study contribute to the database for reference emu hematological and serum enzyme, metabolite, and electrolyte values.

Objective: To characterize the electrocardiogram (ECG) of anesthetized adult emus (Dromaius novaehollandiae)., Animals: Ten clinically healthy adult emus anesthetised for routine physical examination and an electrocardiogram, for both monitoring and investigation into any evidence of cardiac disease., Methods: The ECGs for each emu were obtained in right lateral recumbency, using a modified electrode placement that replicated the standard bipolar leads used in small mammals. Lead II was used for waveform analysis., Results: Median P wave amplitude was 0.55 mV (range: 0.2-0.92 mV) and P wave duration was 0.06 s (0.04-0.09 s). S wave amplitude measured 1.42 mV (0.92-2.12 mV), T wave amplitude 0.67 mV (0.16-0.83 mV) and QRS duration was 0.07 s (0.07-0.12 s). Ninety percent of the QRS complexes were of rS type., Conclusion: Our study provides electrocardiographic baseline data for anesthetized adult emus., (Copyright © 2013 Elsevier B.V. All rights reserved.)

1. An assessment of the efficiency of the acrosome reaction (AR) provides an important predictor of the fertilizing potential of semen and for diagnosis of the causes of infertility. A standardized protocol was therefore developed for initiation of the acrosome reaction in emu spermatozoa in vitro, and the role of CaCl2 or perivitelline membrane (PVM) proteins in determining the outcome of the reaction was investigated. 2. The acrosome reaction (assessed by FITC-PNA) was successfully induced in live spermatozoa by incubation for 2 min in NaCl-TES medium supplemented with 5 mM CaCl2. The maximum response was 32% live acrosome-reacted spermatozoa (LAR) achieved after 10 min incubation. 3. Compared to the outcome with 5 mM CaCl2 or PVM protein alone, the response was significantly better with a combination of PVM protein and CaCl2. 4. A significant variation in the percentage of LAR spermatozoa among individual males was observed. No treatment affected the percentage of dead acrosome-reacted spermatozoa. 5. The results emphasize the important role played by both PVM proteins and Ca(2+) in the in vitro initiation of the acrosome reaction.

Multiflagellate sperm represent a rare defect in mammals and also in the emu where an incidence of only 1% multiflagellate sperm was recorded in semen samples from 15 birds. Biflagellate sperm were the most frequent form of the defect observed, but 3- to 5-tailed sperm were also noted. An association was apparent between multiple tails and macrocephalic sperm, which accounted for 0.6% of multiflagellate sperm. Structural features of the defect were generally similar to those seen in mammals. The duplicated tails shared a single midpiece, which housed supernumerary centriolar complexes, each surrounded by a mitochondrial sheath. A single row of mitochondria separated adjacent centriolar complexes. Elements of the connecting piece (segmented columns, capitellum) appeared normal. The nuclear base appeared flat, staggered, or scalloped depending on the number and depth of additional implantation fossae. Multiflagellate emu sperm of normal head dimensions displayed a widened nuclear base (in the form of an attenuated peripheral nuclear extension) to accommodate the attachment of the additional centriolar complexes. Defective mammalian sperm do not show this modification of the nuclear base as the inherently wider sperm head is able to accommodate the supernumerary centrioles. Although often spiraled around each other, the duplicated principal pieces of the tail were generally separated and free. However, in some cells the proximal parts were collectively bound within the plasmalemma. Multiflagellate sperm appear to have a dual origin with some defective cells originating from incomplete cytokinesis and others as a result of abnormal centriolar duplication., (Copyright © 2012 Elsevier Inc. All rights reserved.)

A protocol for storage of emu semen >6 h has not yet been optimized. The objective was to determine: a) whether sperm quality was adversely affected by sudden exposure to low temperatures (5, 10 and 20 °C) during collection; and b) the effects of three storage temperatures (5, 10 and 20 °C) on survival of emu sperm. In two experiments, each repeated three times on alternate days, ejaculates were diluted 1:1 with precooled (5, 10, or 20°C) UWA-E3 diluent and stored for up to 48 h. Collection temperature, or interaction with either the storage time or storage temperature, had no significant effect on sperm viability, motility, or morphology. Mass Motility Score (2.91-3.27 ± 0.26, mean ± SEM), and percentages of live (72.4-76.2 ± 2.4) and morphologically normal sperm (63.3-64.5 ± 2.3) were comparable among collection temperatures. Conversely, storage temperature and storage time affected (P < 0.05) sperm viability, motility, and morphology. After storage for 48 h, percentages of viable, normal, and motile sperm were higher (P < 0.001) at 5 °C (58.7% ± 1.1, 44.7% ± 1.3, and 50.7% ± 4.9, respectively) and 10 °C (62.6% ± 1.1, 54.1% ± 1.3, and 60.4% ± 4.9) than at 20 °C (27.6% ± 1.1, 20.1% ± 1.3, and 25.9% ± 4.9). Beyond 6 h of storage, the percentage of abnormal sperm was higher (P < 0.001) for storage at 5 °C compared to 10 and 20 °C. After 48 h, bacterial counts were considerably higher at 20 °C compared to 5 and 10 °C (P < 0.001). The pH of stored sperm suspension remained unaffected at 5 and 10 °C, but at 20 °C declined to 6.5 ± 0.03 after 24 h (P < 0.05) and to 6.0 ± 0.03 after 48 h (P < 0.001). We concluded that emu semen could be collected at low ambient temperatures (5-20 °C) without compromising its in vitro storage duration and that semen quality during storage for 48 h was better if it was stored at 10 °C than at 5 or 20 °C., (Copyright © 2012 Elsevier Inc. All rights reserved.)

1. Survival of emu spermatozoa during in vitro storage is not affected by increasing the extracellular [K(+)] to the point where it does not adversely affect spermatozoa function. 2. In three experiments, the effects were studied of [K(+)] in a diluent in the range 12·5-80 mM/l on emu spermatozoa survival for up to 48 h at 5, 10 or 20°C. 3. At the end of the storage period, spermatozoa viability, motility, fertilising ability and morphology were measured. 4. In Experiment 1, spermatozoa viability and morphology were adversely affected after storage (P < 0·001) only in the diluent containing 80 mM/l [K(+)] whereas spermatozoa motility decreased as [K(+)] increased from 12·5 to 80 mM/l. 5. In Experiment 2, during storage at 5°C, the spermatozoa viability was comparable among any of the diluents (standard or modified) but morphology was better (P < 0·001) in all of the modified diluents than in the standard E3 diluent. 6. In Experiment 3, after 48 h of storage in a diluent containing 40 mM/l of [K(+)], the spermatozoa functions were better preserved at 10°C than at 5 or 20°C. 7. It is concluded that a higher than physiological level of potassium can be used in a diluent without detrimental effect on emu spermatozoa survival during 48 h storage and that the best outcome was with storage at 10°C rather than 5 or 20°C.

Three experiments conducted to improve the survival of emu sperm during cryopreservation aimed to: (1) minimize chilling injury during the cooling phase; (2) determine the osmotic effects of dimethylacetamide (DMA), sucrose and trehalose; and (3) investigate the timing and nature of cryoprotectant toxicity. We measured sperm membrane integrity, motility, morphology and egg membrane penetration. In Experiment 1, semen diluted 1:1 with a pre-cooled diluent (5°C) prevented chilling injury. In Experiment 2, semen was diluted with DMA, trehalose or sucrose (300-2400mOsm/L) in deionized water. Only added DMA decreased the percentage of morphologically normal sperm. The percentage of motile sperm was higher with DMA than with the sugars, but membrane intact sperm were comparable amongst all cryoprotectants. As for the osmotic effects, the percentage of membrane intact sperm decreased with 2400mOsm/L and sperm motility decreased with 1200-2400mOsm/L, but sperm morphology was similar at all osmolarities. In Experiment 3, sperm membrane integrity, motility and morphology were comparable at all DMA osmolarities between sperm equilibrated for 0 and 15min, and remained unchanged after removal of DMA. We conclude that: (a) loss of sperm function during the cooling phase can be avoided by using a diluent maintained at 5°C; (b) emu spermatozoa tolerate upto 1400mOsm/L; (c) DMA results in a permanent change in sperm morphology when it is dissolved in deionized water, but does not alter sperm membrane integrity and motility; and (d) equilibration time of sperm with DMA can be less than 10min., (Copyright © 2011 Elsevier B.V. All rights reserved.)

A widely held assumption is that metabolic rate (Ė(met)) during legged locomotion is linked to the mechanics of different gaits and this linkage helps explain the preferred speeds of animals in nature. However, despite several prominent exceptions, Ė(met) of walking and running vertebrates has been nearly uniformly characterized as increasing linearly with speed across all gaits. This description of locomotor energetics does not predict energetically optimal speeds for minimal cost of transport (E(cot)). We tested whether large bipedal ratite birds (emus and ostriches) have gait-specific energetics during walking and running similar to those found in humans. We found that during locomotion, emus showed a curvilinear relationship between Ė(met) and speed during walking, and both emus and ostriches demonstrated an abrupt change in the slope of Ė(met) versus speed at the gait transition with a linear increase during running. Similar to human locomotion, the minimum net E(cot) calculated after subtracting resting metabolism was lower in walking than in running in both species. However, the difference in net E(cot) between walking and running was less than is found in humans because of a greater change in the slope of Ė(met) versus speed at the gait transition, which lowers the cost of running for the avian bipeds. For emus, we also show that animals moving freely overground avoid a range of speeds surrounding the gait-transition speed within which the E(cot) is large. These data suggest that deviations from a linear relation of metabolic rate and speed and variations in transport costs with speed are more widespread than is often assumed, and provide new evidence that locomotor energetics influences the choice of speed in bipedal animals. The low cost of transport for walking is probably ecologically important for emus and ostriches because they spend the majority of their active day walking, and thus the energy used for locomotion is a large part of their daily energy budget.

Osteonecrosis of the femoral head remains a challenging orthopaedic problem. The disease frequently progresses to femoral head collapse, leading to debilitating osteoarthritis in the affected hip(s). Since a major goal of pre-collapse interventions is to forestall the need for hip arthroplasty, it is important that any animal models used to develop or study such interventions also have a natural history of progression to femoral head collapse. The emu (Dromaius novaehollandiae), a large flightless bird native to Australia, consistently progresses to femoral head collapse when osteonecrosis is experimentally induced cryogenically. Full biomechanical characterization of the demands this animal places on its hip is an important consideration in future usage of this model. This study reports in vitro measurement of the contact stress distributions on the emu femoral head during stance phase of the gait cycle, using Fuji pressure-sensitive film. Applied hip loadings were based upon ground reaction forces and hip flexion angles recorded in vivo. The contact stress data showed reasonable homology with the human hip, both in terms of stress magnitude and sites of habitual loading on the femoral head.

The avian embryo has a pair of ductus arteriosi that allow the blood to bypass the pulmonary circulation prior to the initiation of lung ventilation. Our objective was to characterize the factors regulating DA tone during the later stages of development in the emu embryo. We examined in vitro the reactivity of the emu ductus from day 39 through 49 of a 50-day incubation. Steady state tension was not altered by the COX inhibitor indomethacin or the nitric oxide synthase inhibitor L-NAME. However, prostaglandin E(2) (PGE(2)) produced a significant relaxation. Norephinephrine and U-46619 produced strong significant contractions in the emu DA and the adrenergic response matured with development. The contractile response to oxygen matured as the embryo developed with significant oxygen-induced contraction on days 45 and 49, but not on day 39 of incubation. The Kv channel inhibitor 4-aminopyridine induced the contraction of the day 48-49 ductus of similar magnitude as the oxygen-induced contraction. The oxygen-induced contraction was reversed by the reducing agent DTT and the electron transport chain inhibitor rotenone. These results suggest that while the emu DA responds to PGE(2), locally produced PGE(2) are not the important regulators of vessel tone. Additionally, relaxation upon addition of the mitochondria electron transport chain inhibitor rotenone suggests that the mitochondria might be acting as vascular oxygen sensors in this system through the production of reactive oxygen species to stimulate the oxygen-induced contraction in a similar fashion to mammals.

The emu is a large, (bipedal) flightless bird that potentially can be used to study various orthopaedic disorders in which load protection of the experimental limb is a limitation of quadrupedal models. An anatomy-based analysis of normal emu walking gait was undertaken to determine hip contact forces for comparison with human data. Kinematic and kinetic data captured for two laboratory-habituated emus were used to drive the model. Muscle attachment data were obtained by dissection, and bony geometries were obtained by CT scan. Inverse dynamics calculations at all major lower-limb joints were used in conjunction with optimization of muscle forces to determine hip contact forces. Like human walking gait, emu ground reaction forces showed a bimodal distribution over the course of the stance phase. Two-bird averaged maximum hip contact force was approximately 5.5 times body weight, directed nominally axially along the femur. This value is only modestly larger than optimization-based hip contact forces reported in literature for humans. The interspecies similarity in hip contact forces makes the emu a biomechanically attractive animal in which to model loading-dependent human orthopaedic hip disorders.

Most studies examining changes in mechanical performance in animals across size have typically focused on inter-specific comparisons across large size ranges. Scale effects, however, can also have important consequences in vertebrates as they increase in size and mass during ontogeny. The goal of this study was to examine how growth and development in the emu (Dromaius novaehollandiae) hindlimb skeleton reflects the demands placed upon it by ontogenetic changes in locomotor mechanics and body mass. Bone strain patterns in the femur and tibiotarsus (TBT) were related to ontogenetic changes in limb kinematics, ground reaction forces, and ontogenetic scaling patterns of the cross-sectional bone geometry, curvature and mineral ash content over a 4.4-fold increase in leg length and 65-fold increase in mass. Although the distribution of principal and axial strains remained similar in both bones over the ontogenetic size range examined, principal strains on the cranial femur and caudal femur and TBT increased significantly during growth. The ontogenetic increase in principal strains in these bones was likely caused by isometry or only slight positive allometry in bone cross-sectional geometry during growth, while relative limb loading remained similar. The growth-related increase in bone strain magnitude was likely mitigated by increased bone mineralization and decreased curvature. Throughout most of ontogeny, shear strains dominated loading in both bones. This was reflected in the nearly circular cross-sectional geometry of the femur and TBT, suggesting selection for resistance to high torsional loads, as opposed to the more eccentric cross-sectional geometries often associated with the bending common to tetrapods with parasagittal limb orientations, for which in vivo bone strains have typically been measured to date.

Orthopaedic management of femoral head osteonecrosis remains problematic, partly because of inability to systematically compare treatments in an animal model whose natural history parallels the human in terms of progression to femoral head collapse. Recently, it was determined that collapse could be consistently achieved for cryogenically induced osteonecrosis in the emu. Toward delineating the comparative hip joint biomechanics of emus versus humans, for purposes of establishing the emu as a model for human femoral head osteonecrosis, habitual hip joint activity level was quantified for a group of seven healthy adult emus housed in an outdoor research pen typical of those used in emu farming operations. The daily number of steps taken, and the time spent with the hips loaded (standing, or squatting/sitting) versus unloaded (recumbent), were quantified from 24-hour videotape recordings, analyzed by four independent observers. The average number of steps taken per day was 9563, which extrapolates to 1.8 million hip loadings per year, a value that falls in the same general range as seen in normal adult humans. On average, the emus spent 4:05 hours per day idly standing, 2:12 hours squatting/sitting, and 10:44 hours recumbent; they underwent an average of 37 transitions per day between the respective posture/activity states.

In the auditory system, precise encoding of temporal information is critical for sound localization, a task with direct behavioral relevance. Interaural timing differences (ITDs) are computed using axonal delay lines and cellular coincidence detectors in nucleus laminaris (NL). We present morphological and physiological data on the timing circuits in the emu, Dromaius novaehollandiae, and compare these results with those from the barn owl (Tyto alba) and the domestic chick (Gallus gallus). Emu NL was composed of a compact monolayer of bitufted neurons whose two thick primary dendrites were oriented dorsoventrally. They showed a gradient in dendritic length along the presumed tonotopic axis. The NL and nucleus magnocellularis (NM) neurons were strongly immunoreactive for parvalbumin, a calcium-binding protein. Antibodies against synaptic vesicle protein 2 and glutamic acid decarboxlyase revealed that excitatory synapses terminated heavily on the dendritic tufts, while inhibitory terminals were distributed more uniformly. Physiological recordings from brainstem slices demonstrated contralateral delay lines from NM to NL. During whole-cell patch-clamp recordings, NM and NL neurons fired single spikes and were doubly rectifying. NL and NM neurons had input resistances of 30.0 +/- 19.9 Momega and 49.0 +/- 25.6 Momega, respectively, and membrane time constants of 12.8 +/- 3.8 ms and 3.9 +/- 0.2 ms. These results provide further support for the Jeffress model for sound localization in birds. The emu timing circuits showed the ancestral (plesiomorphic) pattern in their anatomy and physiology, while differences in dendritic structure compared to chick and owl may indicate specialization for encoding ITDs at low best frequencies., (Copyright 2006 Wiley-Liss, Inc.)

This study examined ontogenetic patterns of limb loading, bone strains, and relative changes in bone geometry to explore the relationship between in vivo mechanics and size-related changes in the limb skeleton of two vertebrate taxa. Despite maintaining similar relative limb loads during ontogeny, bone strain magnitudes in the goat radius and emu tibiotarsus generally increased. However, while the strain increases in the emu tibiotarsus were mostly insignificant, strains within the radii of adult goats were two to four times greater than in young goats. The disparity between ontogenetic strain increases in these taxa resulted from differences in ontogenetic scaling patterns of the cross-sectional bone geometry. While the cross-sectional and second moments of area scaled with negative allometry in the goat radius, these measures were not significantly different from isometry in the emu tibiotarsus. Although the juveniles of both taxa exhibited lower strains and higher safety factors than the adults, the radii of the young goats were more robust relative to the adult goats than were the tibiotarsi of the young compared with adult emu. Differences in ontogenetic growth and strain patterns in the limb bones examined likely result from different threat avoidance strategies and selection pressures in the juveniles of these two taxa.

Parental investment in eggs and, consequently, in offspring can profoundly influence the phenotype, survival and ultimately evolutionary fitness of an organism. Avian eggs are excellent model systems to examine maternal allocation of energy translated through egg size variation. We used the natural range in emu Dromaius novaehollandiae egg size, from 400 g to >700 g, to examine the influence of maternal investment in eggs on the morphology and physiology of hatchlings. Female emus provisioned larger eggs with a greater absolute amount of energy, nutrients and water in the yolk and albumen. Variation in maternal investment was reflected in differences in hatchling size, which increased isometrically with egg size. Egg size also influenced the physiology of developing emu embryos, such that late-term embryonic metabolic rate was positively correlated with egg size and embryos developing in larger eggs consumed more yolk during development. Large eggs produced hatchlings that were both heavier (yolk-free wet and dry mass) and structurally larger (tibiotarsus and culmen lengths) than hatchlings emerging from smaller eggs. As with many other precocial birds, larger hatchlings also contained more water, which was reflected in a greater blood volume. However, blood osmolality, hemoglobin content and hematocrit did not vary with hatchling mass. Emu maternal investment in offspring, measured by egg size and composition, is significantly correlated with the morphology and physiology of hatchlings and, in turn, may influence the success of these organisms during the first days of the juvenile stage.

Our understanding of avian embryonic cardiovascular regulation has been based on studies in chickens. The present study was undertaken to determine if the patterns established in chickens are generally applicable to the emu, a ratite bird species. We studied cardiovascular physiology over the interval from 60% to 90% of the emu's 50-day incubation period. During this period, embryonic emus exhibit a slight fall in resting heart rate (from 171 beats min(-1) to 154 beats min(-1)) and a doubling of mean arterial pressure (from 1.2 kPa to 2.6 kPa). Exposures to 15% or 10% O(2) initially decreased heart rate during the first period of emu incubation studied [60% of incubation (60%I)] but increased heart rate in the 90%I group. Arterial pressure responded to hypoxia with an initial depression (-1.6 kPa) at 60%I and 70%I but showed no response during the later periods of incubation (80%I and 90%I). In addition, tonic stimulation of both cholinergic and adrenergic (alpha and beta) receptors was present on heart rate at 70%I, with the cholinergic and beta-adrenergic tone increasing in strength by 90%I. Arterial pressure was dependent on a constant beta-adrenergic and constant alpha-adrenergic tone from 60%I to 90%I. A comparison with embryonic white leghorn chickens over a similar window of incubation revealed that emus and white leghorn chickens both possess an adrenergic tone on heart rate and pressure but that only emus possess a cholinergic tone on heart rate. Collectively, these data indicate that the maturation of cardiovascular control systems differs between white leghorn chickens and emus, inviting investigation of additional avian species to determine other patterns.

Among fluctuations of instantaneous heart rate (IHR) in newly hatched chicks, heart rate (HR) oscillation with a mean frequency of 0.7 Hz has been designated as Type II HR variability characterized by low frequency (LF) oscillation [Comp. Biochem. Physiol. Part A 124 (1999) 461]. In response to exposure to lowered ambient temperature (Ta), chick hatchlings raised their HR baseline accompanied with the production or augmentation of Type II HR oscillation, indicating that LF oscillation is a phenomenon relating to thermoregulation [J. Therm. Biol. 26 (2001) 281]. In emu hatchlings that are precocial like chickens, Type II HR oscillation also occurred, but less frequently in comparison with chick hatchlings [Comp. Biochem. Physiol. Part A 131 (2002) 787]. This present experiment was conducted to elucidate how IHR of emu hatchlings responds to changes in Ta. Six hatchlings were measured for IHR and skin temperature (Ts) during a 3-h period when they were exposed to controlled Ta (ca. 35 degrees C), lowered Ta (ca. 15-30 degrees C) and again the controlled Ta for individual 1-h periods. In response to all the cooling and re-warming procedures, HR baseline changed depending upon the intensity of the Ta differences; i.e. large differences of Ta produced large changes in HR. HR fluctuations tended to augment during cooling with a few exceptions, but LF oscillation was not produced. Thus, LF oscillation, which was scarce even at the controlled Ta, could not be used as a thermoregulatory indicator in emus.

In the present study, we determined the distribution of spermatozoa in the perivitelline layer above the germinal disc (GD) of emu and ostrich eggs that had been laid at random intervals after mating. Eggs were opened, the perivitelline layer overlying the GD region was collected and sperm were visualized with 4',6'-diamidino-2-phenylindole under a fluorescence microscope. To map the distribution of sperm, the GD was divided into six areas (A-F), with A being the centre of the GD and F the area furthest from the centre. In both species, more spermatozoa were found in areas B, C and D than in areas A, E and F. More than half the GD spermatozoa were found in areas B, C and D. The pattern of distribution of spermatozoa across the GD depended on the total number of sperm in the GD. In the emu, the pattern was related to delay since last copulation and time of laying, whereas in the ostrich the pattern was related to the month of the season and the sex ratio of the mating system. When the total number of spermatozoa in the GD increased, the number of spermatozoa increased in every area of the GD, but the centre and the outer areas were the least affected. We conclude that sperm numbers are highest in a band immediately around the centre of the GD and then decline with increasing distance from the centre. The low numbers in the centre of the GD may be due to either low attractiveness of the centre for sperm or high attractiveness of the area immediately adjacent to the centre.

Six emu hatchlings were non-invasively measured for electrocardiogram (ECG) from their chest wall using flexible electrodes, and the instantaneous heart rate (IHR) was determined from ECG throughout the first week of post-hatching life. Although the baseline heart rate (HR) was low, approximately 100-200 beats per min (bpm), compared with chick hatchlings, the IHR fluctuated markedly. The fluctuation of IHR comprised HR variability and irregularities that were designated as types I, II and III in chick hatchlings and additional large accelerations distinctive of emu hatchlings. Type I was HR oscillation with a mean frequency of 0.37 Hz (range 0.2-0.7 Hz), i.e. respiratory sinus arrhythmia (RSA). From RSA, breathing frequency in emu hatchlings was estimated to be approximately half of that in chickens. Type II HR oscillation was also found in the emu; the frequency ranged from approximately 0.04 to 0.1 with a mean of 0.06 Hz, and the magnitude tended to be large compared with that of chickens. In addition to type III HRI, which was designated in chickens, large, irregular HR accelerations were characteristic of emu hatchlings. From IHR data, developmental patterns of mean heart rate (MHR) were constructed and plotted on a single graph to inspect the diurnal rhythm of MHR by visual inspection and power spectrum analysis. A circadian rhythm was not clear in the emu hatchlings, in contrast to chick hatchlings, which showed a dominant diurnal rhythm.

Objective: To examine the use of medetomidine for the sedation of captive and wild cassowaries (Casuarius casuarius)., Design: Clinical evaluation after administration of medetomidine by IM injection., Procedure: Nine captive and two wild birds were chemically restrained, with the drug being administered by dart to 10 birds and hand injected to one. Doses of 0.26 to 0.31 mg/kg IM provided light sedation sufficient to allow approach and limited handling. Doses of 0.38 to 0.54 mg/kg IM provided heavy sedation adequate for full clinical examination. Body weights were estimated in six birds and measured in five birds and ranged from 40 to 66 kg. Sternal recumbency occurred in six birds, three in each dose range. In nine birds sedation was reversed with atipamezole at a dose of 15 to 80 mg/kg IM, which produced a return to alertness in 40 to 139 min. Forceful sneezing occurred during recovery in three birds., Conclusion: Medetomidine is a safe and reliable alternative to manual restraint in cassowaries.

Changes in the fertility status of 10 pairs of emus were investigated using egg break-out and numbers of sperm in the perivitelline membrane of the germinal disc (GD) region. After the sexes were separated, sperm in consecutive eggs declined approximately logarithmically at a mean (+/-SEM, n = 10 females) rate of -0.148 +/- 0.021 per log day. Sperm continued to be detected in eggs for 16.5 +/- 1.7 days during which 5.6 +/- 0.6 fertilized eggs were laid. Fertilized eggs that did not contain detectable sperm were laid by five females for a further 2.2 +/- 0.9 days. Based on break-out fertility, the fertile period continued for up to 18.7 +/- 2.1 days, for which the mean number of laid eggs was 6.3 +/- 0.8. An egg with a 50:50 chance of being fertilized would contain 3.5 sperm mm(-2) of GD. Based on the sperm decline model, an egg containing that many sperm would be laid 21 days after the last copulation. In emus that were not separated and allowed to incubate their eggs (n = 3 pairs), the number of sperm in eggs laid before and during incubation declined in a manner similar to that after the last copulation and egg-laying stopped after the females had laid 3.3 +/- 0.3 eggs. After incubation was terminated, females resumed laying within 8.3 +/- 1.2 days and the number of sperm in eggs gradually increased but it did not return to pre-incubation levels. In non-incubating emus (three pairs), the number of sperm in eggs declined as laying progressed, although lit was higher during the period when the first seven eggs were laid than during the period when the rest of eggs were laid (214 +/- 39 v.100 +/- 16 sperm mm(-2) of GD). Sperm numbers varied between successive eggs but a sharp increase followed by a decrease acted as an indicator of recent copulation. There were 8.7 +/- 0.3 such increases per laying period (one per 2.8 +/- 0.2 eggs), a frequency that suggests that emus copulate once weekly. In conclusion, as long as a female emu is supplied with sperm on a weekly basis, she will be fertile but, when copulations stop, she will stop laying soon after. Male fertility appears to fall towards the end of the laying season and it can be affected by egg incubation at any time of the season.

The objective of this study was to establish, for a short-day breeding bird, the male emu, whether the breeding season is principally controlled by changes in photoperiod, and to investigate the endocrine mechanisms involved. Two groups of adult males were subjected to three alternating periods of 150-185 days of 14 h light/day (LD) and 10 h light/day (SD) terminating in a 360-day period of LD or SD. Transfer from LD to SD led to increases in plasma concentrations of luteinizing hormone (LH) and testosterone, after 82 +/- 8 and 73 +/- 3 (SEM) days, and an increase in prolactin concentrations after 115 +/- 12 days. Concentrations of LH and testosterone began to decrease before transfer back to LD, at a time when prolactin concentrations were approaching peak values. Transfer from LD to 360 days of SD resulted in increases in LH and testosterone concentrations, and these terminated after an increase in prolactin concentrations. After transfer from SD to 360 days of LD, plasma concentrations of LH and testosterone began to increase, after delays of 222 +/- 24 and 225 +/- 13 days, and were high at the end of the study, while prolactin values remained depressed throughout. These observations clearly show that seasonal breeding in the emu is directly controlled by changes in photoperiod. The dynamics of the hormonal responses to change of photoperiod suggest that, despite being short-day breeders, the photoregulation of breeding in emus involves mechanisms that are currently accepted for birds, rather than mechanisms that have been proposed for short-day breeding mammals. The initiation of breeding in emus is due to dissipation of photorefractoriness by short days which leads to an increase in the secretion of gonadotrophins to levels that are sufficient to support full reproductive condition. The termination of breeding, while days are still short, is due to the antigonadotrophic action of prolactin which, unusually for birds, increases while the days are still short. In conclusion, breeding activity in male emus is strongly controlled by photoperiod. Emus are short-day breeders, but the central mechanisms that regulate the secretion of reproductive hormones seem to be similar to those previously proposed for long-day breeding birds. The pattern of prolactin secretion in emus suggests an important role for this hormone in the termination of the breeding cycle.

Several different types of cranial kinesis are present within modern birds, enabling them to move (part of) the upper bill relative to the braincase. This movement of the upper bill results from movement of the quadrate and the pterygoid-palatine complex (PPC). The taxon Palaeognathae is characterised by a very distinct PPC and a special type of cranial kinesis (central kinesis) that is very different from that found in the Neognathae. This has led some authors to hypothesise that there is a functional relationship between the morphology of the PPC and the type of cranial kinesis. This hypothesis is tested here by analysing the movement pattern of both the upper bill and the PPC in birds with three different types of cranial kinesis: prokinesis, distal rhynchokinesis and central rhynchokinesis. Movement patterns were determined using a Roentgen stereophotogrammetry method, which made it possible to detect very small displacements (0.5 mm) of bony elements in three dimensions, while the jaw muscles and ligaments remained intact. We found that in all types of kinesis investigated the movements of the quadrate, jugal bars and PPC are similar. Movement of the quadrate is transferred to the upper beak by the jugal bar and the PPC, which moves almost exclusively forwards and backwards, thereby elevating or depressing the upper bill. The differences between the types of kinesis lie only in the position of the point of rotation. These findings indicate that there is no correlation between the specific morphology of the PPC and the type of cranial kinesis. Several other factors, including the external forces applied during food acquisition, may influence the morphology of the PPC. Differences in PPC morphology therefore appear to be the result of different functional demands acting on the system simultaneously but with different strengths, depending on the species.

The emu (Dromaius novaehollandiae) shows potential as a unique animal model for replicating the femoral head collapse process seen in end-stage human osteonecrosis. Since the collapse phenomenon (and interventions to prevent it) involve mechanical processes, it is important to elucidate the similarities and differences of emus versus humans in terms of hip joint biomechanics. A first step for comparison is the intrinsic mechanical properties of the respective bone tissues, as reflected in cortical bone flexural stiffness and strength. In four-point bending, emu cortical bone was found to have an elastic modulus of 13.1 GPa. Its yield stress was determined to be 113 MPa and the ultimate strength was 146 MPa. Emu cortical bone's elastic modulus was similar to that of other avian species, and falls approximately 25% below that of the human (17.3 GPa).

Rate-versus-intensity functions recorded from mammalian auditory-nerve fibers have been shown to form a continuum of shapes, ranging from saturating to straight and correlating well with spontaneous rate and sensitivity. These variations are believed to be a consequence of the interaction between the sensitivity of the hair-cell afferent synapse and the nonlinear, compressive growth of the cochlear amplifier that enhances mechanical vibrations on the basilar membrane. Little is known, however, about the cochlear amplifier in other vertebrate species. Rate-intensity functions were recorded from auditory-nerve fibers in chicks of the emu, a member of the Ratites, a primitive group of flightless birds that have poorly differentiated short and tall hair cells. Recorded data were found to be well fitted by analytical functions which have previously been shown to represent well the shapes of rate-intensity functions in guinea pigs. At the fibers' most sensitive frequencies, rate-intensity functions were almost exclusively of the sloping (80.9%) or straight (18.6%) type. Flat-saturating functions, the most common type in the mammal, represented only about 0.5% of the total in the emu. Below the best frequency of each fiber, the rate-intensity functions tended more towards the flat-saturating type, as is the case in mammals; a similar but weaker trend was seen above best frequency in most fibers, with only a small proportion (18%) showing the reverse trend. The emu rate-intensity functions were accepted as supporting previous evidence for the existence of a cochlear amplifier in birds, the conclusion was drawn further that the nonlinearity observed is probably due to saturation of the hair-cell transduction mechanism.

1. In south-western Australia, male and female emus decrease their food intake when they start breeding in early winter and increase their intake during spring and summer when the breeding season and egg incubation are finished. 2. This annual feeding cycle seems to be under the influence of several environmental factors. Here, we tested the importance of photoperiod using male emus kept in light-controlled rooms with ad libitum access to food and water. 3. Long days increased food intake whereas short days decreased it. Emus fed only during the light hours. 4. Frequency of meals was similar under the 2-day lengths but meal duration was shorter when the emus were on short days than when they were on long days. Thus, day length seemed to affect appetite but not interest in food. 5. Further investigations are needed to test whether these changes in feeding behaviour are a direct consequence of day length or if they are secondary to photoperiod-driven changes in sexual activity.

When exposed to high ambient temperatures, birds defend body temperature by increasing evaporative water loss, via either respiratory or cutaneous water loss. Water deprivation can lead to changes in thermal responses and lower levels of water use for thermoregulation. We have studied the effect of 2-3 wk of water deprivation on the physiological responses of emus during exposure to an ambient temperature of 45 degrees C. Water deprivation led to a delay in the onset of panting (54 vs. 24 min after start of exposure) and to higher body temperatures (38.7 degrees vs. 38.3 degrees C) at the end of exposure to 45 degrees C. After panting was initiated and body temperature stabilised, the water-deprived emus had a lower total evaporative water loss (77 vs. 101 g/h), the same respiratory water loss (70 vs. 72 g/h), and a lower cutaneous water loss (7 vs. 29 g/h) than they did when hydrated. The factor contributing most to the lower total evaporative water loss in the dehydrated emus was a 47% reduction in dry thermal conductance, which led to a decrease in the exogenous environmental heat load and therefore the level of evaporation needed to defend body temperature. We suggest that the decrease in dry thermal conductance follows from the lower level of cutaneous water loss.

Large animal practitioners are called upon to assist producers to prevent and control disease and to manage flocks. The concerns that exist in traditional livestock and poultry management are also applicable to ratites. Adequate ventilation, shelter, and space allocation are the foundations of flock health. Practitioners need to know how to handle ratites in order to perform a through physical examination and collect diagnostic samples.

Seasonal changes in testicular morphology and blood plasma concentrations of LH, testosterone, and prolactin are described for captive male emus in southwestern Australia. Testicular mass and testicular testosterone did not differ between the non-breeding (spring-summer) and the breeding (autumn-winter) seasons. Nevertheless, the testes obtained in the breeding season (May and August) were nearly two fold greater in mass than those collected in the non-breeding season (October and February). The highest testicular concentrations of testosterone were observed in February and lowest in October, while the values during the breeding season were intermediate. The patterns of histological changes in the testes also indicate that emus breed over the autumn-winter months. Tubule diameter was larger in the breeding season than in the non-breeding season, whereas the relative volume of the interstitium was larger in the non-breeding and smaller in the breeding season. Moreover, during the autumn and winter months, plasma LH and testosterone concentrations were high. Outside this period, in spring and summer, the concentrations of these hormones were low. Prolactin concentrations rose around the winter solstice, after the initial increases in plasma LH and testosterone. The end of the breeding season, in early spring, was marked by a gradual decrease in plasma LH concentrations but a rapid fall in testosterone concentrations. Prolactin concentrations continued to increase and peaked near the spring equinox, several weeks after the breeding season ended, and then decreased to reach baseline values by mid-summer. These testicular and endocrine changes are consistent with observations that the emu is a short-day breeder in southwestern Australia. Reproductive activity in the male begins soon after the summer solstice, well in advance of the development of suitable breeding conditions, and is then terminated in spring before food resources become limited by the onset of the dry season.

Similarly sized bipeds and quadrupeds use nearly the same amount of metabolic energy to run, despite dramatic differences in morphology and running mechanics. It has been shown that the rate of metabolic energy use in quadrupedal runners and bipedal hoppers can be predicted from just body weight and the time available to generate force as indicated by the duration of foot-ground contact. We tested whether this link between running mechanics and energetics also applies to running bipeds. We measured rates of energy consumption and times of foot contact for humans (mean body mass 78.88 kg) and five species of birds (mean body mass range 0.13-40.1 kg). We find that most (70-90%) of the increase in metabolic rate with speed in running bipeds can be explained by changes in the time available to generate force. The rate of force generation also explains differences in metabolic rate over the size range of birds measured. However, for a given rate of force generation, birds use on average 1.7 times more metabolic energy than quadrupeds. The rate of energy consumption for a given rate of force generation for humans is intermediate between that of birds and quadrupeds. These results support the idea that the cost of muscular force production determines the energy cost of running and suggest that bipedal runners use more energy for a given rate of force production because they require a greater volume of muscle to support their body weight.