3D printed models are an accurate, cost-effective, and reproducible tool for quantifying terrestrial thermal environments.

Predicting ecological responses to rapid environmental change has become one of the greatest challenges of modern biology. One of the major hurdles in forecasting these responses is accurately quantifying the thermal environments that organisms experience. The distribution of temperatures available within an organism's habitat is typically measured using data loggers called operative temperature models (OTMs) that are designed to mimic certain properties of heat exchange in the focal organism. The gold standard for OTM construction in studies of terrestrial ectotherms has been the use of copper electroforming which creates anatomically accurate models that equilibrate quickly to ambient thermal conditions. However, electroformed models require the use of caustic chemicals, are often brittle, and their production is expensive and time intensive. This has resulted in many researchers resorting to the use of simplified OTMs that can yield substantial measurement errors. 3D printing offers the prospect of robust, easily replicated, morphologically accurate, and cost-effective OTMs that capture the benefits but alleviate the problems associated with electroforming. Here, we validate the use of OTMs that were 3D printed using several materials across eight lizard species of different body sizes and living in habitats ranging from deserts to tropical forests. We show that 3D printed OTMs have low thermal inertia and predict the live animal's equilibration temperature with high accuracy across a wide range of body sizes and microhabitats. Finally, we developed a free online repository and database of 3D scans (https://www.3dotm.org/) to increase the accessibility of this tool to researchers around the world and facilitate ease of production of 3D printed models. 3D printing of OTMs is generalizable to taxa beyond lizards. If widely adopted, this approach promises greater accuracy and reproducibility in studies of terrestrial thermal ecology and should lead to improved forecasts of the biological impacts of climate change.

When Food Fights Back: Skull Morphology and Feeding Behavior of Centipede-Eating Snakes.

Feeding is a complex process that involves an integrated response of multiple functional systems. Animals evolve phenotypic integration of complex morphological traits to covary and maximize performance of feeding behaviors. Specialization, such as feeding on dangerous prey, can further shape the integration of behavior and morphology as traits are expected to evolve and maintain function in parallel. Feeding on centipedes, with their powerful forcipules that pinch and inject venom, has evolved multiple times within snakes, including the genus Tantilla. However, the behavioral and morphological adaptations used to consume this dangerous prey are poorly understood. By studying snakes with varying degrees of dietary specialization, we can test the integration of diet, morphology, and behavior to better understand the evolution of consuming difficult prey. We studied the prey preference and feeding behavior of Tantilla using the flat-headed snake (T. gracilis) and the crowned snake (T. coronata), which differ in the percentage of centipedes in their diet. We then quantified cranial anatomy using geometric morphometric data from CT scans. To test prey preference, we offered multiple types of prey and recorded snake behavior. Both species of snakes showed interest in multiple prey types, but only struck or consumed centipedes. To subdue centipedes, crowned snakes used coiling and holding (envenomation) immediately after striking, while flat-headed snakes used the novel behavior of pausing and holding onto centipedes for a prolonged time prior to the completion of swallowing. Each skull element differed in shape after removing the effects of size, position, and orientation. The rear fang was larger in crowned snakes, but the mechanical advantage of the lower jaw was greater in flat-headed snakes. Our results suggest that the integration of behavioral and morphological adaptations is important for the success of subduing and consuming dangerous prey.

Elevation, oxygen, and the origins of viviparity.

Research focused on understanding the evolutionary factors that shape parity mode evolution among vertebrates have long focused on squamate reptiles (snakes and lizards), which contain all but one of the evolutionary transitions from oviparity to viviparity among extant amniotes. While most hypotheses have focused on the role of cool temperatures in favoring viviparity in thermoregulating snakes and lizards, there is a growing appreciation in the biogeographic literature for the importance of lower oxygen concentrations at high elevations for the evolution of parity mode. However, the physiological mechanisms underlying how hypoxia might reduce fitness, and how viviparity can alleviate this fitness decrement, has not been systematically evaluated. We qualitatively evaluated previous research on reproductive and developmental physiology, and found that (1) hypoxia can negatively affect fitness of squamate embryos, (2) oxygen availability in the circulatory system of adult lizards can be similar or greater than an egg, and (3) gravid females can possess adaptive phenotypic plasticity in response to hypoxia. These findings suggest that the impact of hypoxia on the development and physiology of oviparous and viviparous squamates would be a fruitful area of research for understanding the evolution of viviparity. To that end, we propose an integrative research program for studying hypoxia and the evolution of viviparity in squamates.

A physician associate-led clinic for people with severe mental illness in the United Kingdom.

OBJECTIVE: To evaluate an enhanced physical health clinic led by physician associates (PAs) for patients with severe mental illness. METHODS: A guidance and data collection tool was developed to support and document the outcomes of the PA-led enhanced physical health clinic. RESULTS: The clinic led to diagnoses of diabetes, hyperlipidemia, and hematologic abnormalities. One patient was started on metformin, two patients started a prediabetes program with their general practitioner, one patient started simvastatin, one patient switched from cigarettes to e-cigarettes, and one patient switched from olanzapine to aripiprazole because of metabolic adverse reactions. Three patients intended to contact the National Health Service for cancer screening for which they were eligible but they had not taken up. CONCLUSIONS: PAs can be integrated into a community mental health multidisciplinary team and support the physical health of people with severe mental illness. Mental health trusts should consider roles for PAs in their workforce planning.

Drug-induced Hyperthermic Syndromes in Psychiatry.

Hyperthermia, or extreme elevations in body temperature, can be life-threatening and may be caused by prescription drugs or illegal substances acting at a number of different levels of the neuraxis. Several psychotropic drug classes and combinations have been associated with a classic clinical syndrome of hyperthermia, skeletal muscle hyper-metabolism, rigidity or rhabdomyolysis, autonomic dysfunction and altered mental status ranging from catatonic stupor to coma. It is critical for clinicians to have a high index of suspicion for these relatively uncommon drug-induced adverse effects and to become familiar with their management to prevent serious morbidity and mortality. Although these syndromes look alike, they are triggered by quite different mechanisms, and apart from the need to withdraw or restore potential triggering drugs and provide intensive medical care, specific treatments may vary. Clinical similarities have led to theoretical speculations about common mechanisms and shared genetic predispositions underlying these syndromes, suggesting that there may be a common "thermic stress syndrome" triggered in humans and animal models by a variety of pharmacological or environmental challenges.

Relative fitness of decoy coloration is mediated by habitat type.

Predator-prey interactions can be important drivers of morphological evolution, and antipredator traits in particular. Further, ecological context can be an important factor shaping the evolution of these traits. However, the role of ecological factors such as habitat structure in altering predator-based selection is not well known for antipredator traits such as decoy coloration. We used a combination of a natural history collection survey and a clay model experiment in open- and closed-canopy habitats to study how ecological context alters the fitness benefit of either red or blue decoy coloration in skinks. We found that the development and ecology of red decoy coloration of mole skinks differed substantially from blue tail coloration of other sympatric skink species. Mole skinks do not reach the body size of sympatric species of skinks and retain decoy coloration throughout development. Both patterns of scarring in museum specimens and attacks on plasticine models suggest that red coloration serves as a decoy, attracting attacks to the autotomous tail. While predation rates were similar across habitats, models with red tails were attacked far less frequently in open habitats than models with blue tails, while attack rates were similar in closed habitats. Our results suggest that red decoy coloration in mole skinks could be an adaptation to relatively open-canopy habitats. Our study has important implications for understanding how habitat structure and predator-based selection can alter the evolutionary dynamics of decoy coloration.

Sentence perception in noise by hearing-aid users predicted by syllable-constituent perception and the use of context.

Masked sentence perception by hearing-aid users is strongly correlated with three variables: (1) the ability to hear phonetic details as estimated by the identification of syllable constituents in quiet or in noise; (2) the ability to use situational context that is extrinsic to the speech signal; and (3) the ability to use inherent context provided by the speech signal itself. This approach is called "the syllable-constituent, contextual theory of speech perception" and is supported by the performance of 57 hearing-aid users in the identification of 109 syllable constituents presented in a background of 12-talker babble and the identification of words in naturally spoken sentences presented in the same babble. A simple mathematical model, inspired in large part by Boothroyd and Nittrouer [(1988). J. Acoust. Soc. Am. 84, 101-114] and Fletcher [Allen (1996) J. Acoust. Soc. Am. 99, 1825-1834], predicts sentence perception from listeners' abilities to recognize isolated syllable constituents and to benefit from context. When the identification accuracy of syllable constituents is greater than about 55%, individual differences in context utilization play a minor role in determining the sentence scores. As syllable-constituent scores fall below 55%, individual differences in context utilization play an increasingly greater role in determining sentence scores. Implications for hearing-aid design goals and fitting procedures are discussed.

A Re-evaluation of the Anatomy of the Claustrum in Rodents and Primates-Analyzing the Effect of Pallial Expansion.

The components of the claustrum have been identified by gene expression in mice, but there is still uncertainty about the location of homologous components in primates. To aid interpretation of homologous elements between rodents and primates, we used a current understanding of pallial topology, species-specific telencephalic deformation, and gene expression data. In both rodents and primates, pallial areas maintain conserved topological relationships regardless of relative differences in pallial expansion. The components of the claustrum in primates can, therefore, be identified on the basis of their conserved topological relationships and patterns of gene expression. In rodents, a fairly straight telencephalic long axis runs between the early septopreoptic and amygdalar poles of the pallium. In primates, however, the remarkable dorsal pallial expansion causes this axis to be distorted to form a C shape. This has resulted in a number of errors in the interpretation of the location of claustral components. These errors are likely to have resulted from the unexpected topographical positioning of claustral components due to the bent telencephalic axis. We argue that, once the telencephalic distortion has been accounted for, both rodents and primates have homologous claustral components, and that the topological relationships of these components are conserved regardless of differences in the relative expansion of pallial areas.

Time for Radical Changes in Brain Stem Nomenclature-Applying the Lessons From Developmental Gene Patterns.

The traditional subdivision of the brain stem into midbrain, pons, and medulla oblongata is based purely on the external appearance of the human brain stem. There is an urgent need to update the names of brain stem structures to be consistent with the discovery of rhomobomeric segmentation based on gene expression. The most important mistakes are the belief that the pons occupies the upper half of the hindbrain, the failure to recognize the isthmus as the first segment of the hindbrain, and the mistaken inclusion of diencephalic structures in the midbrain. The new nomenclature will apply to all mammals. This essay recommends a new brain stem nomenclature based on developmental gene expression, progeny analysis, and fate mapping. In addition, we have made comment on the names given to a number of internal brain stem structures and have offered alternatives where necessary.

Spinal Accessory Motor Neurons in the Mouse: A Special Type of Branchial Motor Neuron?

The spinal accessory nerve arises from motor neurons in the upper cervical spinal cord. The axons of these motor neurons exit dorsal to the ligamentum denticulatum and form the spinal accessory nerve. The nerve ascends in the spinal subarachnoid space to enter the posterior cranial fossa through the foramen magnum. The spinal accessory nerve then turns caudally to exit through the jugular foramen alongside the vagus and glossopharyngeal nerves, and then travels to supply the sternomastoid and trapezius muscles in the neck. The unusual course of the spinal accessory nerve has long prompted speculation that it is not a typical spinal motor nerve and that it might represent a caudal remnant of the branchial motor system. Our cell lineage tracing data, combined with images from public databases, show that the spinal accessory motor neurons in the mouse transiently express Phox2b, a transcription factor that is required for development of brain stem branchial motor nuclei. While this is strong prima facie evidence that the spinal accessory motor neurons should be classified as branchial motor, the evolutionary history of these motor neurons in anamniote vertebrates suggests that they may be considered to be an atypical branchial group that possesses both branchial and somatic characteristics. Anat Rec, 302:505-511, 2019. © 2018 Wiley Periodicals, Inc.

Elongation of the CA1 field of the septal hippocampus in ungulates.

It is widely assumed that the hippocampal formation seen in laboratory rodents and in primates is typical of that seen in other mammals. We have tested this assumption by examining sections of brains of 56 mammals from 20 mammalian orders from images on the brainmuseum.org website. We found wide variation in the form of the hippocampal formation, the most extreme examples of which are seen in ungulates, which possess an unusual elongation of the distal CA1 of the septal hippocampus. This phenomenon has not previously been reported. In individual coronal sections of the brains of seven artiodactyl ungulates, the pyramidal layer of CA1 is four times as long as CA2 + CA3. In a perissodactyl ungulate (Burchell's zebra) the distal end of CA1 is so large that it forms a number of folds. A similar but less pronounced CA1 elongation was seen in the brains of 14 carnivores. A modest elongation of CA1 is also present in some other placental mammals, notably the elephant shrew, hyrax, capybara, beaver, and rabbit. The elongation was not present in brains of primates, marsupials, or monotremes. The distal part of CA1 has been shown to play a role in object integration into the spatial map. We hypothesize that the distal CA1 enlargement could serve to enhance the ability to integrate objects into spatial navigation, which would be an advantage for migrating herds of ungulates. We suggest that the remarkable elongation of Q5 CA1 represents a major evolutionary specialization in the ungulates.

Escapees from Rhombomeric Lineage Restriction: Extensive Migration Rostral to the r4/r5 Border of Hox-a3 Expression.

The rhombomeric compartments of the hindbrain are characterized by lineage restriction; cells born in one compartment generally remain there and do not migrate to neighboring rhombomeres. Two well-known exceptions are the substantial migrations of the pontine nuclei and the mammalian facial nucleus. In this study we used Hoxa3-Cre lineage to permanently mark cells that originate in rhombomeres caudal to r4. We found that cells born caudal to the r4/r5 border migrate forwards to a number of different locations in rhombomeres 1-4; the final locations include the interfascicular trigeminal nucleus, the principal trigeminal nucleus, the pontine nuclei, the reticulotegmental nucleus, the ventral nucleus of the lateral lemniscus, and the lateral and medial vestibular nuclei. We suggest that there are numerous exceptions to the principle of rhombomeric lineage restriction that have previously gone unnoticed. Anat Rec, 2017. © 2017 Wiley Periodicals, Inc. Anat Rec, 300:1838-1846, 2017. © 2017 Wiley Periodicals, Inc.

An ontologically consistent MRI-based atlas of the mouse diencephalon.

In topological terms, the diencephalon lies between the hypothalamus and the midbrain. It is made up of three segments, prosomere 1 (pretectum), prosomere 2 (thalamus), and prosomere 3 (the prethalamus). A number of MRI-based atlases of different parts of the mouse brain have already been published, but none of them displays the segments the diencephalon and their component nuclei. In this study we present a new volumetric atlas identifying 89 structures in the diencephalon of the male C57BL/6J 12 week mouse. This atlas is based on an average of MR scans of 18 mouse brains imaged with a 16.4T scanner. This atlas is available for download at www.imaging.org.au/AMBMC. Additionally, we have created an FSL package to enable nonlinear registration of novel data sets to the AMBMC model and subsequent automatic segmentation.

Syllable-constituent perception by hearing-aid users: Common factors in quiet and noise.

The abilities of 59 adult hearing-aid users to hear phonetic details were assessed by measuring their abilities to identify syllable constituents in quiet and in differing levels of noise (12-talker babble) while wearing their aids. The set of sounds consisted of 109 frequently occurring syllable constituents (45 onsets, 28 nuclei, and 36 codas) spoken in varied phonetic contexts by eight talkers. In nominal quiet, a speech-to-noise ratio (SNR) of 40 dB, scores of individual listeners ranged from about 23% to 85% correct. Averaged over the range of SNRs commonly encountered in noisy situations, scores of individual listeners ranged from about 10% to 71% correct. The scores in quiet and in noise were very strongly correlated, R = 0.96. This high correlation implies that common factors play primary roles in the perception of phonetic details in quiet and in noise. Otherwise said, hearing-aid users' problems perceiving phonetic details in noise appear to be tied to their problems perceiving phonetic details in quiet and vice versa.

Mouse Fgf8-Cre-LacZ lineage analysis defines the territory of the postnatal mammalian isthmus.

The isthmus is recognized as the most rostral segment of the hindbrain in non-mammalian vertebrates. In mammalian embryos, transient Fgf8 expression defines the developing isthmic region, lying between the midbrain and the first rhombomere, but there has been uncertainty about the existence of a distinct isthmic segment in postnatal mammals. We attempted to find if the region of early embryonic Fgf8 expression (which is considered to involve the entire extent of the prospective isthmus initially) might help to identify the boundaries of the isthmus in postnatal animals. By creating an Fgf8-Cre-LacZ lineage in mice, we were able to show that Fgf8-Cre reporter expression in postnatal mice is present in the same nuclei that characterize the isthmic region in birds. The 'signature' isthmic structures in birds include the trochlear nucleus, the dorsal raphe nucleus, the microcellular tegmental nuclei, the pedunculotegmental nucleus, the vermis of the cerebellum, rostral parts of the parabrachial complex and locus coeruleus, and the caudal parts of the substantia nigra and VTA. We found that all of these structures were labeled with the Fgf8-Cre reporter in the mouse brain, and we conclude that the isthmus is a distinct segment of the mammalian brain lying caudal to the midbrain and rostral to rhombomere 1 of the hindbrain.

Validation of a Computer-Administered Version of the Digits-in-Noise Test for Hearing Screening in the United States.

BACKGROUND: The sooner people receive treatment for hearing loss (HL), the quicker they are able to recognize speech and to master hearing aid technology. Unfortunately, a majority of people with HL wait until their impairments have progressed from moderate to severe levels before seeking auditory rehabilitation. To increase the number of individuals with HL who pursue and receive auditory rehabilitation, it is necessary to improve methods for identifying and informing these people via widely accessible hearing screening procedures. Screening for HL is the first in a chain of events that must take place to increase the number of patients who enter the hearing health-care system. New methods for hearing screening should be readily accessible through a common medium (e.g., telephone or computer) and should be relatively easy and quick for people to self-administer. PURPOSE: The purpose of this study was to assess a digits-in-noise (DIN) hearing screening test that was delivered via personal computer. RESEARCH DESIGN: Participants completed the Hearing Handicap Inventory for Adults (HHIA) questionnaire, audiometric testing in a sound booth, and computerized DIN testing. During the DIN test, sequences of three spoken digits were presented in noise via headphones at varying signal-to-noise ratios (SNRs). Participants entered each three-digit sequence they heard using an on-screen keypad. STUDY SAMPLE: Forty adults (16 females, 24 males) participated in the study, of whom 20 had normal hearing and 20 had HL (pure-tone average [PTA] thresholds for 0.5, 1, 2, and 4 kHz >25 dB HL). DATA COLLECTION AND ANALYSIS: DIN SNR and PTA data were analyzed and compared for each ear tested. Receiver operating characteristic curves based on these data were plotted. A measure of overall accuracy of a screening test is the area under the receiver operating characteristic curve (AUC). This measures the average true positive rate across false positives at varying DIN SNR cutoffs. Larger values of the AUC indicate, on average, more accurate screening tests. HHIA responses were analyzed and compared to PTA and DIN SNR results using Pearson correlation statistics. RESULTS: HHIA scores were positively correlated with audiometric PTA and DIN SNR results (p < 0.001 for all correlations). For an HL criterion of one or more frequencies from 0.25 to 8 kHz >25 dB HL, the AUC for the DIN test was 0.95. When a criterion of hearling level was set at one or more frequencies from 0.25 to 8 kHz >20 dB HL, the AUC for the DIN test was 0.96. CONCLUSIONS: The computer version of the DIN test demonstrated excellent sensitivity and specificity for our sample of 40 participants. AUC results (≥0.95) suggest that this DIN test administered via computer should be very useful for adult hearing screening.

Developmental gene expression in the mouse clarifies the organization of the claustrum and related endopiriform nuclei.

Studies on gene expression in the developing claustrum of the mouse have clarified the relationships and identity of the claustrum proper and related endopiriform nuclei. The cells of the claustrum primordium express Nr4a2; they are formed in combination with the Nr4a2-labeled subplate cells in the lateral pallium at the site of the future insular cortex. The insular cortex cells, which are born later and which are Nr4a2-negative, migrate through the claustrum toward the pial surface to form layers (2-6a) of the insular cortex. The claustrum is made up of distinct deep (subplate-like) and superficial (principal) parts. The cells of the dorsal endopiriform nucleus (which are also Nr4a2-positive) are formed in the deep part of the claustrum primordium in the lateral pallium, but they migrate ventrally to reach the ventral pallium deep to the piriform cortex at E14.5 in the mouse. On the other hand, the ventral endopiriform nucleus is formed by radially migrating Nr4a2-negative cells in the ventral pallium; it is therefore developmentally distinct from the Nr4a2-postive dorsal endopiriform nucleus, which is a lateral pallial derivative. J. Comp. Neurol. 525:1499-1508, 2017. © 2016 Wiley Periodicals, Inc.

Interspecific Differences in Metabolic Rate and Metabolic Temperature Sensitivity Create Distinct Thermal Ecological Niches in Lizards (Plestiodon).

Three congeneric lizards from the southeastern United States (Plestiodon fasciatus, P. inexpectatus, and P. laticeps) exhibit a unique nested distribution. All three skink species inhabit the US Southeast, but two extend northward to central Ohio (P. fasciatus and P. laticeps) and P. fasciatus extends well into Canada. Distinct interspecific differences in microhabitat selection and behavior are associated with the cooler temperatures of the more Northern ranges. We hypothesized that interspecific differences in metabolic temperature sensitivity locally segregates them across their total range. Resting oxygen consumption was measured at 20°, 25° and 30°C. Plestiodon fasciatus, from the coolest habitats, exhibited greatly elevated oxygen consumption compared to the other species at high ecologically-relevant temperatures (0.10, 0.17 and 0.83 ml O2. g-1. h-1 at 20°, 25° and 30°C, respectively). Yet, P. inexpectatus, from the warmest habitats, exhibited sharply decreased oxygen consumption compared to the other species at lower ecologically-relevant temperatures (0.09, 0.27 and 0.42 ml O2. g-1. h-1 at 20°, 25° and 30°C, respectively). Plestiodon laticeps, from both open and closed microhabitats and intermediate latitudinal range, exhibited oxygen consumptions significantly lower than the other two species (0.057, 0.104 and 0.172 ml O2. g-1. h-1 at 20°, 25° and 30°C, respectively). Overall, Plestiodon showed metabolic temperature sensitivities (Q10s) in the range of 2-3 over the middle of each species' normal temperature range. However, especially P. fasciatus and P. inexpectatus showed highly elevated Q10s (9 to 25) at the extreme ends of their temperature range. While morphologically similar, these skinks are metabolically distinct across the genus' habitat, likely having contributed to their current distribution.

Preventive Dorzolamide-Timolol for Rising Intraocular Pressure During Steep Trendelenburg Position Surgery.

The study purpose was to evaluate preventive use of dorzolamide-timolol ophthalmic solution (Cosopt) during laparoscopic surgery with the patient in steep Trendelenburg (ST) position. Periorbital swelling, venous congestion, and elevated intraocular pressure (IOP) may produce low ocular perfusion. Prompt IOP reduction is important because 30- to 40-minute episodes of acute IOP elevations can result in retinal ganglion cell dysfunction. Dorzolamide-timolol ophthalmic drops reduce IOP and may ameliorate this effect. A double-blind randomized experimental study was conducted to test the effect of dorzolamide-timolol on IOP elevation during laparoscopic surgeries in ST position. Patients were randomly assigned to receive dorzolamide-timolol treatment or balanced salt solution following anesthesia induction. The IOP levels were measured at baseline and 30-minutes intervals throughout surgery. The generalized estimating equations model was used to analyze treatment and time effects and treatment by time interactions. Ninety patients were recruited, with 46 receiving dorzolamide-timolol treatment and 44 receiving balanced salt solution. Statistical analysis revealed significant treatment and time effects and treatment-time interactions on IOP. Patients' IOP was significantly lower in the treatment group than controls (P < .05 to P < .001). Treatment effects were medium to strong. Prophylactic therapy with dorzolamide-timolol significantly reduced IOP of surgical patients during ST positioning.

The organization of spinal motor neurons in a monotreme is consistent with a six-region schema of the mammalian spinal cord.

The motor neurons in the spinal cord of an echidna (Tachyglossus aculeatus) have been mapped in Nissl-stained sections from spinal cord segments defined by spinal nerve anatomy. A medial motor column of motor neurons is found at all spinal cord levels, and a hypaxial column is found at most levels. The organization of the motor neuron clusters in the lateral motor column of the brachial (C5 to T3) and crural (L2 to S3) limb enlargements is very similar to the pattern previously revealed by retrograde tracing in placental mammals, and the motor neuron clusters have been tentatively identified according to the muscle groups they are likely to supply. The region separating the two limb enlargements (T4 to L1) contains preganglionic motor neurons that appear to represent the spinal sympathetic outflow. Immediately caudal to the crural limb enlargement is a short column of preganglionic motor neurons (S3 to S4), which it is believed represents the pelvic parasympathetic outflow. The rostral and caudal ends of the spinal cord contain neither a lateral motor column nor a preganglionic column. Branchial motor neurons (which are believed to supply the sternomastoid and trapezius muscles) are present at the lateral margin of the ventral horn in rostral cervical segments (C2-C4). These same segments contain the phrenic nucleus, which belongs to the hypaxial column. The presence or absence of the main spinal motor neuron columns in the different regions echidna spinal cord (and also in that of other amniote vertebrates) provides a basis for dividing the spinal cord into six main regions - prebrachial, brachial, postbrachial, crural, postcrural and caudal. The considerable biological and functional significance of this subdivision pattern is supported by recent studies on spinal cord hox gene expression in chicks and mice. On the other hand, the familiar 'segments' of the spinal cord are defined only by the anatomy of adjacent vertebrae, and are not demarcated by intrinsic gene expression. The recognition of segments defined by vertebrae (somites) is obviously of great value in defining topography, but the emphasis on such segments obscures the underlying evolutionary reality of a spinal cord comprised of six genetically defined regions. The six-region system can be usefully applied to the spinal cord of any amniote (and probably most anurans), independent of the number of vertebral segments in each part of the spinal column.