Auditory systems of Heteromyidae: functional morphology and evolution of the middle ear.

Middle ears (515) from 26 species of the rodent family Heteromyidae - genera Dipodomys, Microdipodops, Perognathus, and Liomys - were studied both grossly and histologically, for qualitative and quantitative comparisons. Middle ear modifications characteristic of each genus are qualitatively described. Quantitative comparisons are made among the 26 species in the study. Some correlations between middle ear size and other measurements are discussed. The middle ear is an acoustical transformer that for best efficiency must match the impedance of the cochlea to the impedance of the air in the external auditory meatus. It accomplishes this by a pressure increase and a velocity decrease through the combined effects of the lever and areal ratios; however, because the important consideration is a matching of two impedances rather than an absolute pressure increase, the pressure transformer ratio is a less informative measure of the middle ear's efficiency than is the impedance transform ratio. The impedance transformer mechanism is explained (from a morphological point of view), and equations are presented. Dipodomys, Microdipodops, and Perognathus have a theoretical transmission (at the resonant frequency) of 94-100% of the incident acoustical energy; Liomys, 78-80%. The areal ratio of stapes footplate to 2/3 tympanic membrane is remarkably constant among the species, varying only from 0.04 to 0.07: in Dipodomys and Microdipodops this small ratio is due to the very large tympanic membrane; in Perognathus and Liomys it is due to the extremely small stapes footplate. The lever ratio of incus to malleus varies from 0.28 to 0.33 in Dipodpmys and Microdipodops, from 0.37 to 0.46 in Perognathus, and from 0.55 to 0.60 in Liomys. In addition, the middle ear volumes and the morphology of tympanic membrane, ossicles, ligaments, and muscles, all combine to minimize both mass and stiffness. All these data suggest middle ear mechanisms which are very efficient over a broad frequency range. The middle ear modifications found in heteromyids are adaptive in predator avoidance, especially in areas of little natural cover; nevertheless, contrary to expectations, there is no firm relationship between habitat and the extent of these modifications in the 26 species. However, environment did apparently plan an important role in the evolution of the family, and this is discussed.     

Auditory development in the mouse: Structural maturation of the middle ear

The development of middle-ear structures in the mouse was examined in nine groups of pups between 1 and 45 days of age. The area of the tympanic membrane (pars tensa and pars flaccida), the length of the lever arms of the malleus and incus, the surface area of the oval window, and the volume of the bulla all showed systematic changes during neonatal life. The area of the oval window reached maturity first and the lever arms achieved 90% of their adult size on day 11. The tympanic membrane achieved the same criterion on day 18. These data help us further to understand the processes that contribute to the functional ontogeny of the middle ear.     

Solitary is not Asocial: Effects of Social Contact in Kangaroo Rats (Heteromyidae: Dipodomys heermanni)

Social interactions in solitary animals, such as Heermann's kangaroo rat ( Dipodomys heermanni ), have been little studied. In three experiments, the effect of social housing on levels of aggression and maintenance of estrous cycling was studied as part of a captive breeding program for an endangered form of this species. Expt 1 demonstrated that mixed-sex pairs of rats that were neighbors in social housing were no friendlier during staged encounters than unfamiliar pairs also housed socially. In expt 2, rats that had been isolated for several months were transferred to social housing and paired at weekly intervals with an opposite-sex neighbor. Initial encounters were highly aggressive and socialized partners retreated from previously isolated rats more often than vice versa, regardless of sex. By the third week, encounters resembled pairings of fully socialized rats, as in expt 1. Thus, social exposure may produce a general socializing effect rather than promote familiarity between specific individuals. In expt 3, the clear, perforated barriers that separated rats in social housing were replaced with solid, opaque barriers for 6 wk. Estrous cycle length immediately doubled and cycles of most females were disrupted for months. Taken together, the results reveal the influential role of social contact in this solitary species and highlight the need for more such work both for developing comprehensive theories of sociality and for guiding captive propagation efforts.     

Chromosome banding and DNA replication studies on a cell line of Dipodomys merriami

The chromsomes of a cell line of Dipodomys merriami are described in terms of their C-, G- and Q-banding patterns. Studies on the buoyant density of DNA made at different times in the S phase show that the replication of HSα satellite and AT-rich main band DNA occurs preferentially late in the S phase, whereas MS satellite and GC-rich main band DNAs are replicated early in the S phase. Autoradiographic studies of chromosomes labelled early or late in the S phase are used to relate the banding patterns nf particular chromosome regions to the fraction of DNA which they may contain.     

Behavioural syndromes in Merriam's kangaroo rats (Dipodomys merriami): a test of competing hypotheses

Behavioural syndromes, correlations of behaviours conceptually analogous to personalities, have been a topic of recent attention due to their potential to explain trade-offs in behavioural responses, apparently maladaptive behaviour and limits to plasticity. Using Merriam's kangaroo rats (Dipodomys merriami), we assessed the explanatory power and generality of hypothesized syndrome structures derived from the literature and the natural history of the species. Several aspects of functionally distinct behavioural responses of D. merriami were quantified. Syndrome structures were compared using structural equation modelling and model selection procedures. A domain-general behavioural syndrome incorporating cross-functional relationships between measures of boldness, agonistic behaviour, flexibility and food hoarding best explained the data. This pattern suggests that D. merriami behaviours should not be viewed as discrete elements but as components of a multivariate landscape. Our results support arguments that a lack of independence between behaviours may be a general aspect of behavioural phenotypes and suggest that the ability of D. merriami's behaviour to respond to selection may be constrained by underlying connections.     

Patch use by Dipodomys deserti (Rodentia: Heteromyidae): profitability,preference, and depletion dynamics

Summary Granivorous desert rodents of the family Heteromyidae forage nonrandomly among microhabitats that vary in substrate, seed densities, and seed species composition. To explore the hypothesis that microhabitat use is sensitive to seed patch profitability, we quantified effects of seed size (1.96 vs. 5.21 mg/seed) and density (0.4–10.6 seeds/cm²) on Dipodomys deserti harvest rates, which is a measure of profitability when expressed as mg of seed taken per min. By manipulating seed density, we created large-seed and small-seed patches of known relative profitability and exposed D. deserti individuals to pairwise choices in the laboratory and field. We used three treatment classes: 1) large-seed patches that were more profitable than small-seed patches (equal seed densities); 2) large-seed and small-seed patches that were equally profitable (small-seed densities somewhat higher): and 3) large-seed patches that were less profitable than small-seed patches (small-seed densities much higher). Harvest rate increased nearly linearly with seed density, and profitability of large-seed patches was greater than small-seed patches of the same density. Cumulative harvest from a patch increased linearly with residence time up to a plateau; this gain curve indicates that animals move systematically within patches and hence avoid resampling already depleted areas. In the laboratory, animals visited small-seed patches first more often and visited them more frequently when they were more profitable than large-seed patches. When large-seed patches were of greater or equal profitability, large-seed patches were preferred by both measures. The expressed preference for large-seed patches, when animals were presented with equally profitable patches, suggests an underlying preference for large seeds. In the field, animals depleted all patches to a constant low profitability, in accord with qualitative predictions of optimal patch use models. These results suggest that patch preferences by D. deserti are affected by the economics of seed harvest.     

Effects of food value, predation risk, and pilferage on the caching decisions of Dipodomys merriami

Animals that scatter cache their food face a trade-off betweenthe benefits of protecting caches from pilferers and the costsassociated with caching. Placing food into a large number ofwidely spaced caches helps to protect it from pilferage butalso involves costs such as greater exposure to predators. Ipredicted that animals would disperse food into a larger numberof more widely spaced caches when caching (1) a preferred foodversus a less preferred food and (2) under conditions of lowpredation risk versus high predation risk. To test these predictions,I examined the scatter-caching decisions of Merriam's kangaroorats (Dipodomys merriami). D. merriami distributed caches inclumped patterns, regardless of food preference, but they showeda tendency to invest more in a preferred food by distributingcaches more widely. Under the relative safety of the new moon,they did not disperse caches more widely, rather they partitionedthe same amount of food into a larger number of caches thanthey did under the full moon, when predation risk is higher.To examine whether their cache spacing decisions had a significantimpact on the success of cache pilferers, I measured discoveryby pilferers of artificial caches of two food types at differentcaching distances. Results indicate that the cache spacing behaviorof D. merriami functions to protect caches from pilferers, becauseincreased spacing of artificial caches decreased the probabilityof pilferage for both types of food.     

The effect of monaural middle ear destruction on postnatal development of mouse inferior colliculus

This study examined the effect of monaural middle ear destruction on postnatal development of inferior collicular (IC) neurons of the laboratory mouse, Mus musculus. Monaural middle ear destruction was performed on juvenile mice and the density, number and size of IC neurons were determined at different postoperative ages. For electrophysiological study, collicular auditory response properties were always examined four weeks after operation. Monaural middle ear destruction produced larger neurons in the ipsilateral IC (relative to the operated ear) and smaller neurons in the contralateral IC of experimental mice in comparison with IC neurons of control mice. IC neurons of control mice typically had lower minimum thresholds and greater Q10 values than IC neurons of experimental mice. In experimental mice, neurons in the contralateral IC typically had longer latencies and higher minimum thresholds than neurons in the ipsilateral IC. Clear tonotopic organization was only observed for IC neurons of control mice. Possible mechanisms for these different observations are discussed.     

Auditory systems of heteromyidae: Cochlear diversity

Cochleae (125) from 26 species of the rodent family Heteromyidae (genera Dipodomys. Microdipodops, Perognathus, and Liomys) were compared. In Perognathus and Liomys the scala tympani in the apical portion is extremely narrow with a correspondingly minute helicotrema. In Liomys there is no bone separating scala tympani from spiral ganglion in the upper second and entire third turn. In all species studied the zona pectinata of the basilar membrane is enlarged, with a hyaline mass between upper and lower basilar membrane fibers. This zona pectinata hypertrophy is least at the base of the cochlea and greatest in the upper second turn, decreasing again toward the apex. Basilar membrane width increases rapidly in the first turn and then changes only slightly. Except for Liomys, all the heteromyids studied have hypertrophied Hensen's cells with long apical processes supporting and forming an elevated reticular lamina. These Hensen's cells reach their maximum size in the upper second and lower third turns; throughout they rest on inner Claudius' cells rather than the basilar membrane. Relative to naso-occipital length the cochlear specializations are greatest in Microdipodops and least in Liomys just as is the case for middle ear modifications. The morphological data are consistent with the concept that standing wave phenomena may be important in heteromyid cochlear biomechanics. Single unit data of other workers are also consistent with this interpretation. Like middle ear morphology, inner ear morphology appears adapted to low-frequency sensitivity–especially in Dipodomys and Microdipodops.     

The effects of thyroxine on metabolism and water balance in a desert-dwelling rodent, Merriam's kangaroo rat (Dipodomys merriami)

Desert-dwelling mammals such as Merriam's kangaroo rat (Dipodomys merriani) need to conserve both energy and water to survive desert conditions characterized by aridity and low productivity. The thyroid hormone thyroxine increases both basal metabolic rate and urinary water loss in mammals. Increases in basal metabolism and urinary water loss are likely to be detrimental to D. merriami, therefore the regulation of this hormone may be important. To examine the effects of thyroxine in this species, we implanted adult kangaroo rats with pellets designed to release specific doses of thyroxine at a constant rate for 90 days or a placebo pellet. We measured plasma thyroxine concentration, basal metabolic rate, food consumption, urine concentration and water loss in all implanted animals. Thyroxine implants significantly increased both plasma thyroxine and basal metabolic rate in a relatively dose-dependent manner. In response to thyroxine. kangaroo rats increased food consumption only slightly, but this small increase was sufficient to compensate for their elevated metabolic rates. Neither urine concentration nor water loss varied among treatment groups. Thyroxine increased energy expenditure but not water loss in this species.     

The electron-microscopic picture of postnatal development of oligodendroglia.

Ultrastructural studies were carried out on the oligodendroglia in the corpus callosum of rat brain in different periods of the animal's extrauterine development. These investigations showed a possibility of distinguishing in the early stages of ontogenesis three forms of oligodendroblasts differing from one another by electron density and the organelle content of the cytoplasm. The beginning of the production of medullary sheaths was noted as early as the first day after birth and the termination of the morphological phase of this process was recorded on the 8th day of postnatal ontogenesis. Moreover, it was found that on the 17th day of the rat's life the process of differentiation of oligodendroglia in the corpus callosum comes to an end.