Seasonal and daily changes in the behaviour of the adult emu (Dromaius novaehollandiae) under farm conditions.

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.

Visual pigments in a palaeognath bird, the emu Dromaius novaehollandiae: implications for spectral sensitivity and the origin of ultraviolet vision.

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.

Experimental tests of planar strain theory for predicting bone cross-sectional longitudinal and shear strains.

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.

Circulatory changes associated with the closure of the ductus arteriosus in hatching emu (Dromaius novaehollandiae).

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.

A novel transient structure with phylogenetic implications found in ratite spermatids.

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.

Hematological and serum biochemical profile of farm emus (Dromaius novaehollandiae) at the onset of their breeding season.

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.

In vitro initiation of the acrosome reaction in the emu (Dromaius novaehollandiae).

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.

Extending the viability of emu spermatozoa during in vitro storage by manipulation of temperature and diluent potassium concentration.

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.

Gait-specific energetics contributes to economical walking and running in emus and ostriches.

A widely held assumption is that metabolic rate (E(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, E(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 E(met) and speed during walking, and both emus and ostriches demonstrated an abrupt change in the slope of E(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 E(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.

Past and future potential range changes in one of the last large vertebrates of the Australian continent, the emu Dromaius novaehollandiae.

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.

The influence of age and gender on emu (Dromaius novaehollandiae) fat.

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.

Cassowary casques act as thermal windows.

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.

Hip joint contact force in the emu (Dromaius novaehollandiae) during normal level walking.

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.

Comparative analysis of structure and strength of vitelline membrane and physical parameters of yolk of ostrich, emu, and greater rhea eggs.

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.

Habitual hip joint activity level of the penned EMU (Dromaius novaehollandie).

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.

Interaural timing difference circuits in the auditory brainstem of the emu (Dromaius novaehollandiae).

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.

In vivo bone strain and ontogenetic growth patterns in relation to life-history strategies and performance in two vertebrate taxa: goats and emu.

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.

Remote Sensing Derived Fire Frequency, Soil Moisture and Ecosystem Productivity Explain Regional Movements in Emu over Australia.

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.

Fusion Patterns in the Skulls of Modern Archosaurs Reveal That Sutures Are Ambiguous Maturity Indicators for the Dinosauria.

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.