Functional capacity of kangaroo rat hindlimbs: adaptations for locomotor performance.

Many cursorial and large hopping species are extremely efficient locomotors with various morphological adaptations believed to reduce mechanical demand and improve movement efficiency, including elongated distal limb segments. However, despite having elongated limbs, small hoppers such as desert kangaroo rats (Dipodomys deserti) are less efficient locomotors than their larger counterparts, which may be in part due to avoiding predators through explosive jumping movements. Despite potentially conflicting mechanical demands between the two movements, kangaroo rats are both excellent jumpers and attain high hopping speeds, likely due to a specialized hindlimb musculoskeletal morphology. This study combined experimental dissection data with a static analysis of muscle moment generating capacities using a newly developed musculoskeletal model to characterize kangaroo rat hindlimb musculoskeletal architecture and investigate how morphology has evolved to meet hopping and jumping mechanical demands. Hindlimb morphology appears biased towards generating constant moment arms over large joint ranges of motion in this species, which may balance competing requirements by reducing the need for posture and movement specific excitation patterns. The ankle extensors are a major exception to the strong positive relationship exhibited by most muscles between muscle architecture parameters (e.g. Lfibre) and joint moment arms. These muscles appear suited to meeting the high moments required for jumping: the biarticular nature of the ankle extensors is leveraged to reduce MTU strain and create a four-bar linkage that facilitates proximal force transfer. The kangaroo rat hindlimb provides an interesting case study for understanding how morphology balances the sometimes competing demands of hopping and jumping.

Architecture of vasa recta in the renal inner medulla of the desert rodent Dipodomys merriami: potential impact on the urine concentrating mechanism.

We hypothesize that the inner medulla of the kangaroo rat Dipodomys merriami, a desert rodent that concentrates its urine to over 6,000 mosmol/kg H(2)O, provides unique examples of architectural features necessary for production of highly concentrated urine. To investigate this architecture, inner medullary vascular segments in the outer inner medulla were assessed with immunofluorescence and digital reconstructions from tissue sections. Descending vasa recta (DVR) expressing the urea transporter UT-B and the water channel aquaporin 1 lie at the periphery of groups of collecting ducts (CDs) that coalesce in their descent through the inner medulla. Ascending vasa recta (AVR) lie inside and outside groups of CDs. DVR peel away from vascular bundles at a uniform rate as they descend the inner medulla, and feed into networks of AVR that are associated with organized clusters of CDs. These AVR form interstitial nodal spaces, with each space composed of a single CD, two AVR, and one or more ascending thin limbs or prebend segments, an architecture that may lead to solute compartmentation and fluid fluxes essential to the urine concentrating mechanism. Although we have identified several apparent differences, the tubulovascular architecture of the kangaroo rat inner medulla is remarkably similar to that of the Munich Wistar rat at the level of our analyses. More detailed studies are required for identifying interspecies functional differences.

Architecture of kangaroo rat inner medulla: segmentation of descending thin limb of Henle's loop.

We hypothesize that the inner medulla of the kangaroo rat Dipodomys merriami, a desert rodent that concentrates its urine to more than 6,000 mosmol/kgH(2)O water, provides unique examples of architectural features necessary for production of highly concentrated urine. To investigate this architecture, inner medullary nephron segments in the initial 3,000 µm below the outer medulla were assessed with digital reconstructions from physical tissue sections. Descending thin limbs of Henle (DTLs), ascending thin limbs of Henle (ATLs), and collecting ducts (CDs) were identified by immunofluorescence using antibodies that label segment-specific proteins associated with transepithelial water flux (aquaporin 1 and 2, AQP1 and AQP2) and chloride flux (the chloride channel ClC-K1); all tubules and vessels were labeled with wheat germ agglutinin. In the outer 3,000 µm of the inner medulla, AQP1-positive DTLs lie at the periphery of groups of CDs. ATLs lie inside and outside the groups of CDs. Immunohistochemistry and reconstructions of loops that form their bends in the outer 3,000 µm of the inner medulla show that, relative to loop length, the AQP1-positive segment of the kangaroo rat is significantly longer than that of the Munich-Wistar rat. The length of ClC-K1 expression in the prebend region at the terminal end of the descending side of the loop in kangaroo rat is about 50% shorter than that of the Munich-Wistar rat. Tubular fluid of the kangaroo rat DTL may approach osmotic equilibrium with interstitial fluid by water reabsorption along a relatively longer tubule length, compared with Munich-Wistar rat. A relatively shorter-length prebend segment may promote a steeper reabsorptive driving force at the loop bend. These structural features predict functionality that is potentially significant in the production of a high urine osmolality in the kangaroo rat.

Does weather shape rodents? Climate related changes in morphology of two heteromyid species.

Geographical variation in morphometric characters in heteromyid rodents has often correlated with climate gradients. Here, we used the long-term database of rodents trapped in the Sevilleta National Wildlife Refuge in New Mexico, USA to test whether significant annual changes in external morphometric characters are observed in a region with large variations in temperature and precipitation. We looked at the relationships between multiple temperature and precipitation variables and a number of morphological traits (body mass, body, tail, hind leg, and ear length) for two heteromyid rodents, Dipodomys merriami and Perognathus flavescens. Because these rodents can live multiple years in the wild, the climate variables for the year of the capture and the previous 2 years were included in the analyses. Using multiple linear regressions, we found that all of our morphometric traits, with the exception of tail length in D. merriami, had a significant relationship with one or more of the climate variables used. Our results demonstrate that effects of climate change on morphological traits occur over short periods, even in noninsular mammal populations. It is unclear, though, whether these changes are the result of morphological plasticity or natural selection.

Effects of trichloroethylene and perchloroethylene on wild rodents at Edwards Air Force Base, California, USA.

Effects of inhalation of volatilized trichloroethylene (TCE) or perchloroethylene (PCE) were assessed based on the health and population size of wild, burrowing mammals at Edwards Air Force Base (CA, USA). Organic soil-vapor concentrations were measured at three sites with aquifer contamination of TCE or PCE of 5.5 to 77 mg/L and at two uncontaminated reference sites. Population estimates of kangaroo rats (Dipodomys merriami and D. panamintinus) as well as hematology, blood chemistry, and histopathology of kangaroo rats and deer mice (Peromyscus maniculatus) were compared between contaminated and uncontaminated populations. Maximum soil-gas concentrations associated with groundwater contamination were less than 1.5 microl/L of TCE and 0.07 microl/L of PCE. Population estimates of kangaroo rats were similar at contaminated and reference sites. Hematology, blood chemistry, and histopathology of kangaroo rats and deer mice indicated no evidence of health effects caused by exposure. Trichloroethylene or PCE in groundwater and in related soil gas did not appear to reduce the size of small mammal populations or impair the health of individuals.

Developmental and acclimatory contributions to water loss in a desert rodent: investigating the time course of adaptive change.

Understanding the evolution of physiological traits requires considering three nonexclusive mechanisms that underlie phenotypes and cause their change over different time scales: acclimation, developmental plasticity, and natural selection for genetically fixed traits. Physiological adjustments to changes in the desiccating potential of the environment were investigated with one subspecies of common desert rodent, Dipodomys merriami merriami (Merriam's kangaroo rat). We raised young whose parents originated from environments that differ in both temperature and humidity. These young were raised under either desiccating or water-abundant conditions, and their water loss was measured at a series of temperatures to determine the effect developmental conditions have on resistance to desiccation. We then determined the contribution of acclimation to desiccation resistance by keeping the differentially raised young in conditions opposite to those during their development and again measuring water loss. We found that developmental plasticity and acclimation can completely account for the existing intraspecific variability in desiccation resistance under certain conditions. In fact, developmental and acclimatory changes can equal genetically based differences of the populations. This phenotypic plasticity can operate relatively quickly and therefore may attenuate the actions of natural selection. Understanding the extent and nature of such flexibility is critical to our understanding intraspecific variability and the consequences of changing climate.

Natural space-use patterns and hippocampal size in kangaroo rats.

The size of the hippocampus, a forebrain structure that processes spatial information, correlates with the need to relocate food caches by passerine birds and with sex-specific patterns of space use in microtine rodents. The influences on hippocampal anatomy of sexual selection within species, and natural selection between species, have not yet been studied in concert, however. Here we report that natural space-use patterns predict hippocampal size within and between two species of kangaroo rats (Dipodomys). Differences in foraging behavior suggest that Merriam's kangaroo rats (D. merriami) require better spatial abilities than bannertail kangaroo rats (D. spectabilis). Sex-specific differences in mating strategy suggest that males of both species require more spatial ability than females. As predicted, hippocampal size (relative to brain size) is larger in Merriam's than in bannertail kangaroo rats, and males have larger hippocampi than females in both species. Males of a third species (D. ordii) also have smaller hippocampi than Merriam's kangaroo rat males, despite being similar to Merriam's in brain and body size. These results suggest that both natural and sexual selection affect the relative size and perhaps function of mammalian hippocampi. They also reassert that measures of functional subunits of the brain reveal more about brain evolution than measures of total brain size.

[Karyological characteristics of cell sublines of the kidney of the kangaroo rat and of skin fibroblasts of the Indian muntjac]. / Kariologicheskaia kharakteristika kletochnykh sublinii pochki kengurovoi krysy i fibroblastov kozhi indiiskogo muntzhaka.

A study was made of the karyotypic structure of sublines derived from the kangaroo rat's kidney (NBL-3) and skin fibroblasts of the Indian muntjac, available in the cell culture bank of the Institute of Cytology Acad. Sci. USSR. A comparative karyologic analysis was made of subline NBL-3 both contaminated with mycoplasma (NBLK) and decontaminated with antibiotics (NBLD). Authentic differences in cell distribution according to chromosome number in NBLK and NBLD variants were shown. Modal numbers of chromosomes are 11 and 17, respectively. The modal number for the Indian muntjac cell subline (MT) is 9. 60-80% of the cells had an identical karyotype (the main structural variant of the karyotype is MSVK). Using the G-banding technique, all the MSVK variants were shown to display constant karyotypes. In NBLK there are 5 pairs of homologous chromosomes and one metacentric. In NBLD, the number of homologous chromosomes increased in all the groups (hypotriploid karyotype). In subline MT 3 homologous chromosomes were found in groups I and IV, 2 in group III in addition to one X-chromosome. A comparison with the Indian muntjac karyotype showed the absence of marker chromosomes in MT. The analysis of additional SVK shows that the deviations from MSVK are caused mostly by changes in the number of homologous chromosomes within the groups. A study of the frequency of deviations in chromosome numbers observed in the groups from MSVK showed that different chromosomes were involved in karyotypic changes in the same way in the "low-chromosome" variants of NBLK and MT, and in different ways in NBLD. A comparison of the "premycoplasmic" variants of line NBL-3 with NBLK shows no differences in the parameters studied.(ABSTRACT TRUNCATED AT 250 WORDS)

Ultrastructure of pinealocytes of the kangaroo rat (Dipodomys ordi).

The ultrastructure of the pinealocytes of the wild-captured ord kangaroo rat (Dipodomys ordi) was examined. A homogeneous population of pinealocytes was present in the pineal gland of the kangaroo rat. The Golgi apparatus, granular endoplasmic reticulum, mitochondria, lysosomes, dense-core vesicles, vacuoles containing a flocculent material and lipid droplets were consistent components of the pinealocyte cytoplasm, whereas infrequently-observed organelles included centrioles, multivesicular bodies, subsurface cisternae, "synaptic" ribbons and cilia. The number of dense-core vesicles was relatively high and dense-core vesicles and vacuoles containing a flocculent material were present in the same cell. Although it has been recently suggested that two different secretory processes, i.e., neurosecretory-like (Golgi apparatus - dense-core vesicles) and ependymal-like (granular endoplasmic reticulum - vacuoles containing a flocculent material) may be involved in different regulatory mechanisms in the pinealocytes, the definitive answer to this is still far from clear. Therefore, the pineal gland of the kangaroo rat appears to be a good model for the study of the potential relationship between these two secretory processes, especially in respect to seasonal changes.

Morphological study of the desert heteromyid kidney with emphasis on the genus Perognathus.

The renal morphology of three species of desert dwelling Perognathus rodents were compared to Dipodomys and two species of sympatric cricetid rodents. Perognathus has a highly adapted unipolar kidney capable of urine concentration up to 7,500 mOsm/KgH2O. Two major modifications were observed in these kidneys. (1) There is elongation of both the inner and outer medulla. When the thickness of the regions in P. penicillatus are factored by kidney weight it is found that the outer medulla is 3.8 and the inner medulla is 2.2 times greater in length than the extensively studied Dipodomys merriami. (2) There is great variance in glomerular size with approximately 20% of the glomeruli being very large and located in the juxta medullary region. These glomeruli are 1.5--1.6 times greater in diameter than the more numerous superficial glomeruli. These structural modifications are quantitated and the functional implications are discussed particularly in relation to urine concentrating ability in these desert inhabitants.

An anatomical study of the brains of Dipodomys (Mammalia: Rodentia: Heteromyidae).

Whole brains and Nissl-stained serial sections of 3 species of kangaroo rat (Dipodomys spectabilis, D. ordii, D. merriami) were compared for interspecific differences. Neuroanatomical variations were conservative in nature. Examination of the cerebellum revealed possible differences in locomotion and ecology. Dipodomys ordii exhibited more neurological evidence for increased specialization of the hindlimbs, while D. merriami had the least differentiated cerebellar surface, thereby suggesting more stereotyped movements of the hindlimbs. Dipodomys spectabilis had abilities that lie somewhere between those of D. ordii and D. merriami. Stereological analysis demonstrated little difference in the percent of total brain volume comprising the telencephalon, diencephalon, cerebellum, and brain stem; and it revealed the relative position of the cranial nerve nuclei.

Fine structure of the heterochromatin of the kangaroo rat Dipidomys ordii, and examination of the possible role of actin and myosin in heterochromatin condensation.

Biochemical studies have suggested that some actin and myosin may be present in the nucleus. This raises the possibility that heterochromatin condensation might be the result of an actin-myosin rigour type complex. Since ATP dissociates actin and myosin, this possibility could be examined by determining the effect of ATP on heterochromatin condensation. Thin-section electron microscopy showed large amounts of condensed constitutive heterochromatin in the kidney nuclei and somewhat less in the liver nuclei of the kangaroo rat, Dipidomys ordii. Surprisingly, there were some nuclei in the brain which contained no condensed heterochromatin despite the fact that this genome is composed of 50% satellite DNA. Although washing kidney nuclei with solutions of 10 mM Tris-ATP caused marked decondensation of the heterochromatin, when they were washed with Mg-ATP the heterochromatin was more condensed than in the controls. This suggests the decondensation by Tris-ATP is due to its ability to chelate divalent cations and provides no support for condensation of heterochromatin being the result of myosin-actin interaction. Despite being decondensed, the chromatin fibres of heterochromatin were distinct from those of euchromatin. The heterochromatin formed rod-like 19-5 nm fibres, the euchromatin formed random coils of 11-0-nm fibres.

Projections of the trapezoid body and the superior olivary complex of the Kangaroo rat (Dipodomys merriami).

Glass micropipettes filled with 2 M sodium cyanide were used to physiologically locate and iontophoretically damage the nucleus of the trapezoid body (NTB), the medial superior olive (MSO), and the lateral superior olive (LSO). Mechanical lesions were made in the trapezoid body as it leaves the cochlear nuclei. After a 3- to 10-day survival time the projections and terminal degeneration were traced with the Fink-Heimer and Nauta-Gygax stains. The ventral cochlear nucleus (VCN) projects via the trapezoid body to ipsilateral LSO, ipsilateral preolivary nuclei, ipsilateral lateral and a contralateral medial dendritic fields of MSO, and contralateral NTB; there is also a small ipsilateral projection to the ventral nucleus of the lateral lemniscus (VNLL) and the central nucleus of the inferior colliculus (CNIC). Some trapezoid body fibers ascend via the contralateral lateral lemniscus to VNLL, DNLL (dorsal nucleus of the lateral lemniscus), and CNIC. There is no projection from the ventral cochlear nucleus to the ipsilateral NTB and contralateral preolivary nuclei. All portions of NTB project ipsilaterally to LSO (ventral NTB to dorsomedial LSO, dorsal NTB to ventral LSO) and to the retro-olivary nucleus. In two animals with NTB lesions there is also degeneration in the ventromedial portion of the ipsilateral facial nucleus. NTB projects contralaterally by way of the stria of Monakow to the pyramidal and molecular cell layers of the dorsal cochlear nucleus (DCN). The NTB does not project ipsilaterally to MSO, preolivary nuclei, VNLL, DNLL and CNIC. Contralaterally there are no projections to any of the nuclei of the auditory pathway except the DCN. Most MSO projections are ipsilateral. The densest goes by way of the lateral lemniscus to the lateral aspect of the ipsilateral CNIC, terminating throughout its dorsoventral axis. MSO also projects bilaterally to the pyramidal and molecular cell layers of dorsal cochlear nucleus (DCN), and ipsilaterally to the ventral portion of the motor nucleus of V and to the facial nucleus. MSO does not project ipsilaterally to the LSO, NTB, preolivary, VCN and retro-olivary nuclei. On the contralateral side, all structures except the DCN are free of projection patterns from axons originating in the MSO. LSO projects bilaterally to the central and ventral portions of CNIC and to the nuclei of the lateral lemnisci, and ipsilaterally to the large and small spherical cell areas of anterior ventral cochlear nucleus (AVCN) and to all portions of DCN. The LSO does not project ipsilaterally to the NTB, MSO, preolivary and retro-olivary nuclei. On the side opposite, this nucleus does not project to NTB, MSO, retro-olive, VCN, preolivary and LSO. For all lesions regardless of the site, there is no degeneration found rostral to the CNIC. The medial geniculate body or other structures in the diencephalon or cortex are free of any fields of terminal degeneration.