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PubMed2005

The interaction of a new motor skill and an old one: H-reflex conditioning and locomotion in rats.

New and old motor skills can interfere with each other or interact in other ways. Because each skill entails a distributed pattern of activity-dependent plasticity, investigation of their interactions is facilitated by simple models. In a well characterized model of simple learning, rats and monkeys gradually change the size of the H-reflex, the electrical analog of the spinal stretch reflex. This study evaluates in normal rats the interactions of this new skill of H-reflex conditioning with the old well established skill of overground locomotion. In rats in which the soleus H-reflex elicited in the conditioning protocol (i.e., the conditioning H-reflex) had been decreased by down-conditioning, the H-reflexes elicited during the stance and swing phases of locomotion (i.e., the locomotor H-reflexes) were also smaller. Similarly, in rats in which the conditioning H-reflex had been increased by up-conditioning, the locomotor H-reflexes were also larger. Soleus H-reflex conditioning did not affect the duration, length, or right/left symmetry of the step cycle. However, the conditioned change in the stance H-reflex was positively correlated with change in the amplitude of the soleus locomotor burst, and the correlation was consistent with current estimates of the contribution of primary afferent input to the burst. Although H-reflex conditioning and locomotion did not interfere with each other, H-reflex conditioning did affect how locomotion was produced: it changed soleus burst amplitude and may have induced compensatory changes in the activity of other muscles. These results illustrate and clarify the subtlety and complexity of skill interactions. They also suggest that H-reflex conditioning might be used to improve the abnormal locomotion produced by spinal cord injury or other disorders of supraspinal control.

Animals↗
PubMed2005

Degradation of head direction cell activity during inverted locomotion.

Head direction (HD) cells in the rat limbic system carry information about the direction the head is pointing in the horizontal plane. Most previous studies of HD functioning have used animals locomoting in an upright position or ascending/descending a vertical wall. In the present study, we recorded HD cell activity from the anterodorsal thalamic nucleus while the animal was locomoting in an upside-down orientation. Rats performed a shuttle-box task requiring them to climb a vertical wall and locomote across the ceiling of the apparatus while inverted to reach an adjoining wall before ascending into the reward compartment. The apparatus was oriented toward the preferred direction of the recorded cell, or the 180 degrees opposite direction. When the animal was traversing the vertical walls of the apparatus, the HD cells remained directionally tuned as if the walls were an extension of the floor. When the animal was locomoting inverted on the ceiling, however, cells showed a dramatic change in activity. Nearly one-half (47%) of the recorded cells exhibited no directional specificity during inverted locomotion, despite showing robust directional tuning on the walls before and after inversion. The remaining cells showed significantly degraded measures of directional tuning and random shifts of the preferred direction relative to the floor condition while the animal was inverted. It has previously been suggested that the HD system uses head angular velocity signals from the vestibular system to maintain a consistent representation of allocentric direction. These findings suggest that being in an inverted position causes a distortion of the vestibular signal controlling the HD system.

Action Potentials↗
PubMed2006

Coordination and modulation of locomotion pattern generators in Drosophila larvae: effects of altered biogenic amine levels by the tyramine beta hydroxlyase mutation.

Forward locomotion of Drosophila melanogaster larvae is composed of rhythmic waves of contractions that are thought to be produced by segmentally organized central pattern generators. We present a systematic description of spike activity patterns during locomotive contraction waves in semi-intact wild-type and mutant larval preparations. We have shown previously that Tbetah(nM18) mutants, with altered levels of octopamine and tyramine, have a locomotion deficit. By recording en passant from the segmental nerves, we investigated the coordination of the neuronal activity driving contraction waves of the abdominal body-wall muscles. Rhythmic bursts of activity that occurred concurrently with locomotive waves were frequently observed in wild-type larvae but were rarely seen in Tbetah(nM18) mutants. These centrally generated patterned activities were eliminated in the distal stumps of both wild-type and Tbetah(nM18) larvae after severing the segmental nerve from the CNS. Patterned activities persisted in the proximal stumps deprived of sensory feedback from the periphery. Simultaneous recordings demonstrated a delay in the bursting activity between different segments, with greater delay for segments that were farther apart. In contrast, bilateral recordings within a single segment revealed a well synchronized activity pattern in nerves innervating each hemisegment in both wild-type and Tbetah(nM18) larvae. Significantly, rhythmic patterns of bursts and waves could be evoked in Tbetah(nM18) mutants by head or tail stimulation despite their highly irregular spontaneous activities. These observations suggest a role of the biogenic amines in the initiation and modulation of motor pattern generation. The technique presented here can be readily extended to examine the locomotion motor program of other mutants.

Abdominal Wall↗
PubMed2002

The requirement of presynaptic metabotropic glutamate receptors for the maintenance of locomotion.

Spinal circuits known as central pattern generators maintain vertebrate locomotion. In the lamprey, the contralaterally alternating ventral root activity that defines this behavior is driven by ipsilateral glutamatergic excitation (Buchanan and Grillner, 1987) coupled with crossed glycinergic inhibition (Buchanan, 1982; Alford and Williams, 1989). These mechanisms are distributed throughout the spinal cord. Glutamatergic excitatory synapses activate AMPA and NMDA receptors known to be necessary for the maintenance of the locomotor rhythm. Less is known of the role and location of metabotropic glutamate receptors (mGluRs), although group I mGluRs enhance transmitter release at giant synapses in the lamprey spinal cord, whereas group II/III receptors may inhibit release. In this study we show that group I mGluR antagonists block fictive locomotion, a neural correlate of locomotion, by acting at the presynaptic terminal. Under physiological conditions, synaptically released glutamate activates presynaptic group I mGluRs (autoreceptors) during the repetitive activation of glutamatergic terminals. The resulting rise in [Ca2+]i caused by the release from presynaptic intracellular stores is coincident with an enhancement of synaptic transmission. Thus, blocking mGluRs reduces glutamate release during the repetitive activity that is characteristic of locomotion, leading to the arrest of locomotor activity. We found the effects of group I mGluRs on locomotion to be inconsistent with a postsynaptic effect on the central pattern generator. Consequently, the activation of metabotropic glutamate autoreceptors is necessary to maintain rhythmic motor output. Our results demonstrate the role of presynaptic mGluRs in the physiological control of movement for the first time.

Animals↗
PubMed2000

Estimation of the energy costs of locomotion in the Iberian pig (Sus mediterraneus).

The energy cost of locomotion of four Iberian pigs was measured in two experiments conducted when the animals averaged 41.3 (SE 0.1) kg (first experiment) and 84.1 (SE 0.1) kg (second experiment). The heat production of the pigs was determined when standing or walking at a speed of 0.555 m/s on a treadmill enclosed in a confinement-type respiration chamber, on different slopes (-10.5, 0, and +10.5% in the first experiment, and -5.25, 0 and +10.5% in the second experiment). The energy costs of locomotion, estimated from the coefficients of linear regressions of heat production per kg body weight (BW) on distance travelled, were in the first experiment 2.99, 3.31 and 5.88 J/kg BW per m for -10.5, 0, and +10.5% inclines respectively, and 2.56, 2.84 and 7.13 J/kg BW per m for -5.25, 0 and +10.5% inclines respectively, in the second experiment. The net energy cost of locomotion on the level appeared to be independent of live weight, attaining a value of 2.98 J/kg BW per m. Also, it was found that within experiments the net energy cost of walking on negative slopes was similar to that for locomotion on the level, indicating that no energy was recovered on vertical descent. Mean values were 3.11 and 2.72 kJ/kg BW per m for the light and heavy pigs respectively. The energy cost of raising 1 kg BW one vertical metre was found to be 27.1 J/kg BW per m in the first experiment and 40.0 J/kg BW per m in the second experiment. Correspondingly, the calculated efficiency for upslope locomotion appeared to decline with increasing BW, resulting in average values of 36.2 and 24.5%.

Animals↗
PubMed2003

Treadmill locomotion in the intact and spinal mouse.

Because the genetic characteristics of several inbred strains of mice are well identified, their use is becoming increasingly popular in spinal cord injury research. In this context, it appears particularly important to document adequately motor patterns, such as locomotion in normal mice, to establish some baseline values of locomotor characteristics. It also seems crucial to determine the extent to which mice can express a locomotor pattern after a complete spinal transection to establish a baseline on which one can evaluate the effects of treatments after spinal injury. Therefore, we have used conventional techniques to document the kinematics of treadmill locomotion in intact mice (n = 11) and in mice with a complete section of the spinal cord at T8 (n = 12). The results show that the kinematics and EMG of adult normal mice can be adequately monitored with such conventional equipment and that mice can re-express hindlimb locomotion within 14 d after spinalization, without any pharmacological treatments. The angular excursions of the hip, knee, and ankle are similar to those of the intact mice, although the joints are sometimes more flexed. After spinal cord transection, out-of-phase alternation between the homologous limbs recovered, whereas the timing between homolateral limbs was completely lost. This remarkable ability of mice to express hindlimb locomotion after a complete spinalization should be taken into account in the evaluation of various procedures aimed at promoting the functional recovery of locomotion after spinal lesions.

Animals↗
PubMed2005

[Comparative investigation of influence of injections of chlorpromazine and 5,6-dihydroxytryptamine on locomotion, defensive reactions of snall helix lucorum and excitability of command neurons in long-term sensitization].

Comparative analysis of the action of chlorpromazine (CPZ) and neurotoxin 5,6-dihydroxytryptamine (5,6-DHT) on defensive reactions and locomotion of grape snail and elaboration of long-term sensitization (LTS), was carried out. Long-term (chronic) injection of chlorpromazine led to significant increasing of a pneumostome closing time and to changing of motor behaviour towards decrease of the velocity of the locomotion. Daily injections of 5,6-DHT in small doses within a week were accompanied by the gradual decrease of the velocity of snails locomotion, which was kept for a week. Similar effect was observed in injection of neurotoxin (30 mgs/kg). Injections of CPZ prevents elaboration of LTS, as well as injections of 5,6-DHT. After the action of CPZ, LTS, LTS followed by CPZ, and also during elaboration of LTS after injection of CPZ, the velocity of locomotion directly depended on the length of leg. During elaboration of LTS after injection of 5,6-DHT, such dependency is not retained. Electrophysiological study revealed that chronic injections of CPZ led to depolarizing shift of membrane potential and decrease of the threshold of action potential generation in command neurons as after injection of neurotoxin 5,6-DHT. Therefore, the action of neuroleptic drug CPZ on the defensive behaviour, locomotion of grape snail and electrical characteristics of identifying neurons is comparable with the action of toxic analogue of serotonin.

5,6-Dihydroxytryptamine↗
PubMed1991

Factors that determine the magnitude and time course of human H-reflexes in locomotion.

The soleus H-reflex amplitude is deeply modulated during locomotion in humans (Capaday and Stein, 1986). Moreover, at a constant stimulus intensity, the slope of the relationship between the amplitude of the soleus H-reflex and the background electromyogram (EMG) changes with different locomotor tasks (Capaday and Stein, 1987a). Two further aspects are studied here. First, we recorded the reflex during overlapping speeds of walking (2.0-7.5 km/hr) and running (5-9 km/hr) to determine whether the speed, the motor output, or the form of locomotion was most important in setting the slope of this relationship between H-reflex and background EMG. Second, we determined the time course of change in the H-reflex amplitude and the possible site of action for the reflex depression during the transition from standing to walking. The primary determinant of the slope was found to be the form of locomotion. The differences between running and walking could not be explained entirely by either movement speed or motor output. For walking, the slope varied inversely with the speed and the motor output of locomotion. This compensation in slope as a function of motor output may prevent saturation of the motoneuron pool. The appropriate reflex amplitudes for a particular locomotor pattern are activated rapidly and completely within a reaction time, and simultaneously with the activation of muscle activity for the initiation of walking. Mechanisms for the rapid change seen during the initiation of locomotion most likely act presynaptically on the muscle spindle afferents. The time course and magnitude of this change are correlated with the activity of the tibialis anterior muscle.

Electromyography↗
PubMed1987

Diencephalic regulation of respiration and arterial pressure during actual and fictive locomotion in cat.

The purpose of this study was to examine by experimentation the hypothesis that the respiratory and circulatory responses during exercise are attributable to command signals that emanate from the suprapontine brain. We studied the relations between locomotion (exercise) and phrenic nerve activity and arterial pressure in cats that walked or ran on a treadmill and in animals during fictive locomotion, i.e., locomotor activity in motor nerves to legs. Anesthetized cats with intact brains and unanesthetized decorticated cats were used. All preparations exhibited spontaneous actual and fictive locomotion. Electrical stimulation or microinjection of picrotoxin, a GABA antagonist, of the subthalamic locomotor areas always caused locomotion to develop. Phrenic nerve activity and arterial pressure increased in proportion to the level of locomotor activity despite control or ablation of feedback signals from chemoreceptors and vagal receptors. Similar relations were measured during fictive locomotion despite the absence of muscular contraction and limb movement and the lack of change in metabolic rate. These findings provide experimental support for the central command hypothesis for the genesis of the respiratory hyperpnea and increased cardiovascular function that occur during exercise. We believe that the command signals emanate from the subthalamic locomotor area of the diencephalon.

Animals↗
PubMed1993

Electrical systems for improving locomotion after incomplete spinal cord injury: an assessment.

Simple systems for electrical stimulation (1-4 channels) with either surface, percutaneous, or implanted electrodes during locomotion were assessed in 10 subjects who had chronic, incomplete spinal cord injury (SCI). On average, the speed of locomotion was increased by 4 m/min independently of the subject's speed of locomotion without stimulation (0-50 m/min) while oxygen consumption was reduced somewhat. These simple systems can provide practical help, particularly for incomplete SCI subjects who can stand but are lacking or have very limited ability to walk. Further improvement in locomotion requires stabilization and reduction in the duration of the stance phase of locomotion.

Adult↗
PubMed1996

Visuomotor regulation of locomotion.

Vision is obviously important for the control of goal-directed locomotion. This profound influence can be best revealed by interfering with the visual pathways or by altering the visual inputs normally expected to occur during locomotion. For instance, the use of reversed optic flow or reversing prisms can induce significant locomotor changes or lead to powerful illusions, such as the sensation of backward walking while actually walking forward. To better understand the many ways by which visual inputs could actually influence locomotion, an overview of some of the background concepts on the generation, initiation, and control of locomotion is given and, when pertinent, mechanisms by which vision could interact with locomotion at different control levels of the central nervous system are postulated.

Animals↗
PubMed1998

Phasic firing time locked to cocaine self-infusion and locomotion: dissociable firing patterns of single nucleus accumbens neurons in the rat.

The activity of single nucleus accumbens (NAcc) neurons of rats was extracellularly recorded during intravenous cocaine self-administration sessions (0.7 mg/kg per infusion, fixed ratio 1). We reported previously that NAcc neurons showed a change, usually a decrease, in firing rate during the first 1 min after the cocaine-reinforced lever press. This postpress change was followed by a progressive reversal of that change, which began within the first 2 min after the press and was not complete until the last 1 min before the next lever press (termed the change + progressive reversal firing pattern). In the present study we documented a regular pattern of locomotion that occurred in parallel with the change + progressive reversal firing pattern. This observation suggested that discharges time locked to locomotion may determine the change + progressive reversal firing pattern. However, 55% of the neurons failed to show firing time locked to locomotion that could have contributed to the change + progressive reversal firing pattern. Moreover, for all neurons, the change + progressive reversal firing pattern was apparent even if the calculation of firing rate excluded all periods of locomotion. The present data showed that the change + progressive reversal firing pattern is not solely attributable to phasic changes in firing time locked to the execution of locomotion. The change + progressive reversal firing pattern closely mirrors changes in drug level and dopamine overflow observed by previous researchers and may thus be a component of the neurophysiological mechanism by which drug level regulates drug-taking behavior during an ongoing self-administration session.

Action Potentials↗
PubMed1997

Suppression of bleb formation, locomotion, and polarity of Walker carcinosarcoma cells by hypertonic media correlates with cell volume reduction but not with changes in the F-actin content.

The putative role of cellular or solvent volume in protrusive activity and locomotion has been investigated in blebbing Walker carcinosarcoma cells using hypertonic media. Blebbing, locomotion, and cell polarity are completely suppressed by 0.2 M sorbitol. The response occurs in two steps. In a first step, i.e. within 10 sec after the addition of sorbitol, blebbing and locomotion are inhibited and this is associated with an average cell volume reduction by 17% (corresponding to a reduction in solvent volume by 38%). It clearly precedes suppression of cell polarity (pre-existing protrusions, tail) occurring in a second step within 5 to 10 min after addition of sorbitol without additional reduction in the cell or solvent volume. The relative amount of F-actin does not correlate with the decrease in cell volume, suppression of blebbing, locomotion, and cell polarity. A significant decrease in the relative amount of F-actin is found only at volume reductions which are higher than those required to completely suppress blebbing, locomotion, and cell polarity. F-actin staining occurs preferentially along the cell membrane in isotonic as well as in hypertonic media. The results are best compatible with the hypothesis that hydrostatic pressure rather than actin polymerization at the front is the direct force driving the membrane forward during bleb formation. Cells with lamellipodia show a similar response to hypertonic media, suggesting that basically similar mechanisms may operate in both forms of protrusions.

Actins↗
PubMed1985

Phorbol myristate acetate (PMA) suppresses polarization and locomotion and alters F-actin content of Walker carcinosarcoma cells.

This study demonstrates a novel feature of PMA, its ability to suppress chemokinetic polarization and locomotion of tumor cells. Walker carcinosarcoma cells exhibit two distinct types of polarization and locomotion, i.e. spontaneous polarization characterized by ruffles at the front and stimulated polarization and locomotion in response to the microtubule-disassembling agents colchicine, vinblastine and nocodazole, which are characterized by blebbing at the front. The tumor promotor phorbol myristate acetate (PMA), but not phorbol, was found to suppress both types of polarization and random locomotion at concentrations between 10(-8) and 10(-6)M. The effect of 10(-6)M PMA was virtually complete within 5 min. Inhibition of locomotion was due to both a reduction in the speed of migrating cells and the proportion of migrating cells. Changes in shape and chemokinesis of Walker carcinosarcoma cells were associated with alterations in the relative amount and the topographical distribution of F-actin as determined by NBD-phallacidin binding. Suppression by PMA was associated with loss of the polar topographical distribution of F-actin visualized by NBD-phallacidin binding. In the presence of PMA, the relative amount of F-actin was higher than in unstimulated controls and lower in cells exposed to microtubule-disassembling agents.

Actin Cytoskeleton↗
PubMed1989

Diacylglycerols and the protein kinase inhibitor H-7 suppress cell polarity and locomotion of Walker 256 carcinosarcoma cells.

We show that diacylglycerols, like phorbol myristate acetate (PMA), suppress cell polarity and locomotor activity of Walker carcinosarcoma cells in a dose-dependent fashion in vitro. OAG and diC8 show significant activity at concentrations above 3 x 10(-5) M. The inhibitory effect on locomotion is due to a reduction in the proportion of locomoting cells rather than gradual lowering of the speed of individual cells. Measurement of protein kinase C (PKC) activity in isolated fractions showed a substantial reduction of the total cellular PKC activity and of the activity in the cytosolic fraction following incubation of cells with 10(-8) M PMA for 30 min. In contrast, the total and relative PKC activity associated with the membrane fraction was increased by PMA. The effect of H-7, an inhibitor of PKC as well as of cAMP-dependent kinase, has been tested. H-7 suppressed cell polarity of "unstimulated" control cells (ID50 = 6.5 microM H-7), colchicine-stimulated cells (ID50 = 92 microM H-7) or cells treated with both PMA and colchicine (ID50 = 15 microM H-7), in a dose-dependent fashion. The locomotor activity of the cells was also suppressed. LTB4 had no clearcut activity in this system. Our findings suggest that diacylglycerols and H-7 are of interest as physiological or pharmacological stop signals for tumor-cell locomotion. Contrary to our expectations, PMA and diacylglycerols vs. H-7 did not produce opposing or antagonistic effects on cell polarity and locomotion. This similarity may be due to down-regulation of PKC by PMA and inhibition of PKC by H-7. However, the mechanisms underlying these novel effects of diacylglycerols and of H-7 on cell polarity and locomotion may be even more complex; they require further studies.

1-(5-Isoquinolinesulfonyl)-2-Methylpiperazine↗
PubMed1995

Selective responses (actin polymerization, shape changes, locomotion, pinocytosis) to the PKC inhibitor Ro 31-8220 suggest that PKC discriminately regulates functions of human blood lymphocytes.

The results suggest that protein kinase C (PKC) plays a pivotal role in the control of F-actin levels, locomotion, pinocytosis, and cell shape in lymphocytes. The PKC inhibitor Ro 31-8220 elicits a high proportion of polarized (ED50 = 1.5 x 10(-6) M) and locomoting cells and reduces the relative amount of F-actin (by 29% at 10(-5) M) in initially resting cells. Phorbol myristate acetate (PMA) counterbalances the polarizing effect of Ro 31-8220. This indicates that the spherical shape and the F-actin content of resting cells are maintained by constitutive PKC activity. PMA-induced increases in fluid pinocytosis, F-actin content, and formation of nonpolar cells with surface protrusion are suppressed by Ro 31-8220 (IC50 = 2-4 x 10(-7) M). Spherical cells and, at higher concentrations (ED50 = 3.3 x 10(-6) M), polarized cells are formed instead. As a result, lymphocyte function switches from fluid pinocytosis to cell polarity and locomotion. The data indicate that PKC is instrumental in selectively switching lymphocyte function between resting state, locomotor activity, and fluid pinocytosis. Ro 31-8220 is extremely potent in stimulating lymphocyte polarity and locomotion (B and T cells). It acts faster and/or produces a higher proportion of polarized lymphocytes than other available agonists. It may thus be used as a tool in further experiments requiring locomoting lymphocytes.

Actins↗
PubMed1993

Ras-transformed cells express both CD44 and RHAMM hyaluronan receptors: only RHAMM is essential for hyaluronan-promoted locomotion.

Hyaluronan (HA) is an important regulator of cell locomotion. We show that ras-transformed cells, termed 245 cells, respond to HA with an increase in random locomotion. We show that two HA receptors, RHAMM (receptor for hyaluronan-mediated motility) and CD44, are present on these ras-transformed fibroblasts. RHAMM is expressed as a 58-kDa protein and is distributed primarily as patches over lamellae. CD44 occurs largely as an 85- to 90-kDa protein that is distributed more or less evenly over the cell surface with small amounts concentrated at the tips of lamellae. CD44 and RHAMM both bind biotinylated HA in a transblot assay, indicating that they are both potential fibroblast HA receptors. CD44 binds approximately five times more HA than RHAMM as determined by densitometric analysis of transblots, indicating that this protein is the major HA receptor on fibroblasts. We assessed the role of these receptors in mediating the stimulatory effects of HA on cell motility by using antibody neutralization. Several antibodies to CD44 were used that inhibit HA/CD44 interactions. None of these had an effect on locomotory responses to HA, indicating that CD44 is not directly involved in mediating locomotion in response to HA on ras-transformed cells. In contrast, antibodies specific to RHAMM completely inhibited locomotion, indicating that RHAMM is the primary mediator of HA-promoted locomotion of ras-transformed cells.

Animals↗
PubMed1993

Origin of the central entrainment of respiration by locomotion facilitated by MK 801 in the decerebrate rabbit.

In order to establish the origin of the central coupling between locomotion and respiration which operates in freely moving mammals during galloping, we sought experimental conditions that readily lead to such a coupling in decerebrate and curarised rabbit preparations. In such preparations, stimulation of the mesencephalic locomotor region (MLR) evokes locomotor activities, recorded from hindlimb muscle nerves, that are rarely totally coordinated with phrenic inspiratory activity. However, low doses (0.2 mg/kg i.v.) of MK 801, a non-competitive NMDA antagonist which has been shown to increase the activity of the spinal locomotion generators (Fenaux et al. 1991), dramatically enhanced this coupling during MLR stimulation in most experiments: 1/1 coupling was dominant but 2/1 and 3/1 couplings (i.e. two or three locomotor cycles per respiratory cycle) were also obtained. Compared with spontaneous respiratory activity, which was apneustic under these conditions, the respiratory period was drastically decreased during coupling. However, a further transection of the spinal cord at the C6 or C7 level, which isolated the spinal locomotion generators from the supraspinal levels, totally suppressed this reduction of the inspiratory period during MLR stimulation in the presence of MK 801. In experiments where locomotor activity was simultaneously recorded at forelimb and hindlimb levels, the 1/1 evoked locomotor-respiratory coupling remained after the lumbar cord had been isolated by L1 spinal transection. The present data do show that intact spinal mechanisms are required for entrainment to occur. They suggest either tha a common supraspinal drive cannot entrain locomotion and respiration when being depressed, or that respiration is entrained at the locomotor rate by the spinal locomotion generators.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗