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1.
The weakly electric fish Apteronotus leptorhynchus produces wave-like electric organ discharges distinguished by a high degree of regularity. Transient amplitude and frequency modulations (“chirps”) can be evoked in males by stimulation with the electric field of a conspecific. During these interactions, the males examined in this study produced six types of chirps, including two novel ones. Stimulation of a test fish with a conspecific at various distances showed that two electrically interacting fish must be within 10 cm of each other to evoke chirping behavior in the neighboring fish. The chirp rate of all but one chirp type elicited by the neighboring fish was found to be negatively correlated with the absolute value of the frequency difference between the two interacting fish, but independent of the sign of this difference. Correlation analysis of the instantaneous rates of chirp occurrence revealed two modes of interactions characterized by reciprocal stimulation and reciprocal inhibition. Further analysis of the temporal relationship between the chirps generated by the two fish during electric interactions showed that the chirps generated by one individual follow the chirps of the other with a short latency of approximately 500–1000 ms. We hypothesize that this “echo response” serves a communicatory function.  相似文献   

2.
Summary The electric organ discharge (EOD) potential was mapped on the skin and midplane of several Apteronotus leptorhynchus. The frequency components of the EOD on the surface of the fish have extremely stable amplitude and phase. However, the waveform varies considerably with different positions on the body surface. Peaks and zero crossings of the potential propagate along the fish's body, and there is no point where the potential is always zero. The EOD differs significantly from a sinusoid over at least one third of the body and tail. A qualitative comparison between fish showed that each individual had a unique spatiotemporal pattern of the EOD potential on its body.The potential waveforms have been assembled into high temporal and spatial resolution maps which show the dynamics of the EOD. Animation sequences and Macintosh software are available by anonymous ftp (mordor.cns.caltech.edu; cd/pub/ElectricFish).We interpret the EOD maps in terms of ramifications on electric organ control and electroreception. The electrocytes comprising the electric organ do not all fire in unison, indicating that the command pathway is not synchronized overall. The maps suggest that electroreceptors in different regions fulfill different computational roles in electroreception. Receptor mechanisms may exist to make use of the phase information or harmonic content of the EOD, so that both spatial and temporal patterns could contribute information useful for electrolocation and communication.Abbreviations EOD electric organ discharge - EO electric organ - CV coefficient of variance  相似文献   

3.
The African electric fish Gymnarchus niloticus rhythmically emits electric organ discharges (EODs) for communication and navigation. The EODs are generated by the electric organ in the tail in response to the command signals from the medullary pacemaker complex, which consists of a pacemaker nucleus (PN), two lateral relay nuclei (LRN) and a medial relay nucleus (MRN). The premotor structure and its modulatory influences on the pacemaker complex have been investigated in this paper. A bilateral prepacemaker nucleus (PPn) was found in the area of the dorsal posterior nucleus (DP) of the thalamus by retrograde labeling from the PN. No retrogradely labeled neurons outside the pacemaker complex were found after tracer injection into the LRN or MRN. Accordingly, anterogradely labeled terminal fibers from PPn neurons were found only in the PN. Iontophoresis of l-glutamate into the region of the PPn induced EOD interruptions. Despite the exclusive projection of the PPn neurons to the PN, extracellular and intracellular recordings showed that PN neurons continue their firing while MRN neurons ceased their firing during EOD interruption. This mode of EOD interruption differs from those found in any other weakly electric fishes in which EOD cessation mechanisms have been known.  相似文献   

4.
Aperonotus leptorhynchus (Gymnotiformes) produces wave-like electric organ discharges distinguished by a high degree of constancy. Transient frequency and amplitude modulations of these discharges occur both spontaneously and during social interactions, which can be mimicked by external electrical stimulation. The so-called chirps can be divided into four different types. Independent of the type of chirp produced under spontaneous conditions, the fish generate only significant numbers of type-2 chirps under evoked conditions. The rate of production of chirps of this type is largely determined by the frequency relative to the fish's frequency and signal intensity. Frequencies of + 10 Hz of the fish's own discharge frequency most effectively elicit chirps. Type-2 chirps can also be evoked through stimulation at or near the higher harmonic frequencies of the fish's frequency, but the chirp rate decreases with increasing number of the higher harmonic component. Over a certain range, the rate of production of type-2 chirps increases with increasing stimulus intensity. At very high intensities the generation of type-2 chirps is accompanied by the production of a novel type of electrical signal ("abrupt frequency rise") characterized by a frequency increase of approximately 20 Hz and high repetition rates of roughly 10 s(-1). We hypothesize that the different types of electric modulations subserve different behavioral functions.  相似文献   

5.
Serotonin modulates agonistic and reproductive behavior across vertebrate species. 5HT1A and 5HT1B receptors mediate many serotonergic effects on social behavior, but other receptors, including 5HT2 receptors, may also contribute. We investigated serotonergic regulation of electrocommunication signals in the weakly electric fish Apteronotus leptorhynchus. During social interactions, these fish modulate their electric organ discharges (EODs) to produce signals known as chirps. Males chirp more than females and produce two chirp types. Males produce high-frequency chirps as courtship signals; whereas both sexes produce low-frequency chirps during same-sex interactions. Serotonergic innervation of the prepacemaker nucleus, which controls chirping, is more robust in females than males. Serotonin inhibits chirping and may contribute to sexual dimorphism and individual variation in chirping. We elicited chirps with EOD playbacks and pharmacologically manipulated serotonin receptors to determine which receptors regulated chirping. We also asked whether serotonin receptor activation generally modulated chirping or more specifically targeted particular chirp types. Agonists and antagonists of 5HT1B/1D receptors (CP-94253 and GR-125743) did not affect chirping. The 5HT1A receptor agonist 8OH-DPAT specifically increased production of high-frequency chirps. The 5HT2 receptor agonist DOI decreased chirping. Receptor antagonists (WAY-100635 and MDL-11939) opposed the effects of their corresponding agonists. These results suggest that serotonergic inhibition of chirping may be mediated by 5HT2 receptors, but that serotonergic activation of 5HT1A receptors specifically increases the production of high-frequency chirps. The enhancement of chirping by 5HT1A receptors may result from interactions with cortisol and/or arginine vasotocin, which similarly enhance chirping and are influenced by 5HT1A activity in other systems.  相似文献   

6.
I recorded the electric organ discharges (EODs) of 331 immature Brachyhypopomus pinnicaudatus 6–88 mm long. Larvae produced head-positive pulses 1.3 ms long at 7 mm (6 days) and added a second, small head-negative phase at 12 mm. Both phases shortened duration and increased amplitude during growth. Relative to the whole EOD, the negative phase increased duration until 22 mm and amplitude until 37 mm. Fish above 37 mm produced a “symmetric” EOD like that of adult females. I stained cleared fish with Sudan black, or fluorescently labeled serial sections with anti-desmin (electric organ) or anti-myosin (muscle). From day 6 onward, a single electric organ was found at the ventral margin of the hypaxial muscle. Electrocytes were initially cylindrical, overlapping, and stalk-less, but later shortened along the rostrocaudal axis, separated into rows, and formed caudal stalks. This differentiation started in the posterior electric organ in 12-mm fish and was complete in the anterior region of fish with “symmetric” EODs. The lack of a distinct “larval” electric organ in this pulse-type species weakens the hypothesis that all gymnotiforms develop both a temporary (larval) and a permanent (adult) electric organ. Accepted: 1 March 1997  相似文献   

7.
Brown ghost knife fish, Apteronotus leptorhynchus, produce sexually dimorphic, androgen-sensitive electrocommunication signals termed chirps. The androgen regulation of chirping has been studied previously by administering exogenous androgens to females and measuring the chirping response to artificial electrical signals. The present study examined the production of chirps during dyadic interactions of fish and correlated chirp rate with endogenous levels of one particular androgen, 11-ketotestosterone (11KT). Eight males and four females were exposed to short-term (5-min) interactions in both same-sex and opposite-sex dyads. Twenty-four hours after all behavioral tests, fish were bled for determination of plasma 11KT levels. Males and females differed in both their production of chirps and their ability to elicit chirps from other fish: males chirped about 20-30 times more often than females and elicited 2-4 times as many chirps as females. Among males, chirp rate was correlated positively with plasma 11KT, electric organ discharge frequency, and body size. Combined with results from experimental manipulation of androgen levels, these results support the hypothesis that endogenous 11KT levels influence electrocommunication behavior during interactions between two male fish.  相似文献   

8.
When animals are under stress, glucocorticoids commonly inhibit adult neurogenesis by acting through glucocorticoid receptors (GRs). However, in some cases, conditions that elevate glucocorticoids promote adult neurogenesis, and the role of glucocorticoid receptors in these circumstances is not well understood. We examined the involvement of GRs in social enhancement of brain cell addition and aggressive signaling in electric fish, Apteronotus leptorhynchus. In this species, long-term social interaction simultaneously elevates plasma cortisol, enhances brain cell addition and increases production of aggressive electrocommunication signals (“chirps”). We implanted isolated and paired fish with capsules containing nothing (controls) or the GR antagonist, RU486, recorded chirp production and locomotion for 7 d, and measured the density of newborn cells in the periventricular zone. Compared to isolated controls, paired controls showed elevated chirping in two phases: much higher chirp rates in the first 5 h and moderately higher nocturnal rates thereafter. Treating paired fish with RU486 reduced chirp rates in both phases to those of isolated fish, demonstrating that GR activation is crucial for socially induced chirping. Neither RU486 nor social interaction affected locomotion. RU486 treatment to paired fish had a partial effect on cell addition: paired RU486 fish had less cell addition than paired control fish but more than isolated fish. This suggests that cortisol activation of GRs contributes to social enhancement of cell addition but works in parallel with another GR-independent mechanism. RU486 also reduced cell addition in isolated fish, indicating that GRs participate in the regulation of cell addition even when cortisol levels are low.  相似文献   

9.
The neural circuit that controls the electric organ discharge (EOD) of the brown ghost knifefish (Apteronotus leptorhynchus) contains two spontaneous oscillators. Both pacemaker neurons in the medulla and electromotor neurons (EMNs) in the spinal cord fire spontaneously at frequencies of 500–1000 Hz to control the EOD. These neurons continue to fire in vitro at frequencies that are highly correlated with in vivo EOD frequency. Previous studies used channel blocking drugs to pharmacologically characterize ionic currents that control high‐frequency firing in pacemaker neurons. The goal of the present study was to use similar techniques to investigate ionic currents in EMNs, the other type of spontaneously active neuron in the electromotor circuit. As in pacemaker neurons, high‐frequency firing of EMNs was regulated primarily by tetrodotoxin‐sensitive sodium currents and by potassium currents that were sensitive to 4‐aminopyridine and κA‐conotoxin SIVA, but resistant to tetraethylammonium. EMNs, however, differed from pacemaker neurons in their sensitivity to some channel blocking drugs. Alpha‐dendrotoxin, which blocks a subset of Kv1 potassium channels, increased firing rates in EMNs, but not pacemaker neurons; and the sodium channel blocker μO‐conotoxin MrVIA, which reduced firing rates of pacemaker neurons, had no effect on EMNs. These results suggest that similar, but not identical, ionic currents regulate high‐frequency firing in EMNs and pacemaker neurons. The differences in the ionic currents expressed in pacemaker neurons and EMNs might be related to differences in the morphology, connectivity, or function of these two cell types. © 2005 Wiley Periodicals, Inc. J Neurobiol, 2006  相似文献   

10.
1. Sternopygus macrurus were collected in Venezuela during the period of gonadal recrudescence in early or late dry season. Electric organ discharge (EOD) frequencies were recorded, blood samples were taken for analysis of steroid titers, and gonads were taken for determination of reproductive condition. 2. Mean EOD frequencies were significantly lower in males than in females in all samples. EOD frequency was inversely correlated with body length in males in late, but not early, dry season, and these parameters were never correlated in females. 3. Plasma levels of testosterone (T) and 11-ketotestosterone (11-KT), but not estradiol-17 beta (E2), were inversely correlated with EOD frequency in males. No 11-KT was observed in plasma of females, and plasma levels of T and E2 in females were comparable to those of males. Neither T nor E2 were correlated with EOD frequency in females. 4. Testes collected in late dry season were more mature than those from early dry season; androgen levels and EOD frequency were correlated with testicular maturity. Ovaries collected in early dry season were immature, while those from late dry season were more mature. There was no relationship between EOD frequency and stage of ovarian development. 5. These results suggest that plasma androgens modulate EOD frequency in males during the reproductive season and that plasma E2 has little relationship to EOD frequency in either sex.  相似文献   

11.
The ability to orient to and track moving electrolocation targets was assessed in normal Apteronotus leptorhynchus and in those with unilateral lesions of the nucleus praeeminentialis dorsalis.
  1. Each fish was trained to hover between two vertical metal rods and track their movement. Two aspects of this behavior were measured: a) the hovering position of the fish relative to stationary rods; b) the latency between the onset of rod motion and the fish's tracking response. Control fish hovered midway between stationary rods, while lesioned fish hovered closer to the rod ipsilateral to the lesion. Response latency varied negatively with rod diameter in both sets of fish, and lesioned fish exhibited shorter latencies than control fish. While the response latencies of control fish were shortest when their starting position was midway between the rods, lesioned animals' latencies were shortest when they hovered closer to the rod ipsilateral to their lesion.
  2. Control fish responded to the approach of a single metal ball to either side of the body with nearly equal latencies and fish-to-object distances. After lesioning, response latency increased and fish-to-object distance decreased for approaches to the side ipsilateral to the lesion; opposite changes occurred for contralateral approaches.
  相似文献   

12.
Summary A classical conditioning paradigm was used to test the ability of Sternopygus macrurus to detect EOD-like stimuli (sine waves) of different frequencies. The behavioral tuning curves were quite close in shape to tuning curves based on single-unit recordings of T units, although the sensitivity at all frequencies was much greater. The behavioral curves showed notches of greatly reduced sensitivity when the test frequency was equal to, or twice the EOD frequency. The EOD of each of the fish was eliminated by lesioning the medullary pacemaker nucleus, and the fish were retested. The resulting tuning curves were nearly the same in shape as those of the EOD-intact individuals, but the PMN-lesioned fish showed an overall reduction of sensitivity of 30 dB. The EOD appears to enhance sensitivity by placing the summed stimulus (test stimulus + fish's EOD) at an amplitude where T units are maximally sensitive to small temporal modulations in the fish's own EOD. Peripheral tuning appears to limit the ability of males to detect the EOD of females, since these are, on average, an octave higher in frequency than the male EOD, while the peak sensitivity of the male occurs 5–10 Hz above its own EOD frequency.Abbreviations EOD electric organ discharge - PMN pacemaker nucleus - BF best frequency - DF difference frequency  相似文献   

13.
There is a sexual dimorphism in the frequency of the quasi-sinusoidal electric organ discharge (EOD) of Sternopygus macrurus, with males, on average, an octave lower. EODs are detected by tuberous electroreceptor organs, which exhibit V-shaped frequency tuning with maximal sensitivity near the fish's own EOD frequency. This would seem to limit the ability of a fish to detect the EODs of opposite-sex conspecifics. However, electroreceptor tuning has always been based on single-frequency stimulation, while actual EOD detection involves the addition of a conspecific EOD to the fish's own. In the present study, recordings were made from single electroreceptive units while the fish were stimulated with pairs of sine waves: one (S1) representing the fish's own EOD added to a second (S2) representing a conspecific EOD. T unit response was easily predicted by assuming that the electroreceptor acts as a linear filter in series with a threshold-sensitive spike initiator. P unit response was more complex, and unexpectedly high sensitivity was found for frequencies of S2 well displaced from the fish's EOD frequency. For both P and T units, detection thresholds for S2 were much lower when added to S1, than when presented alone.  相似文献   

14.
The two closely related gymnotiform fishes, Apteronotus and Eigenmannia, share many similar communication and electrolocation behaviors that require modulation of the frequency of their electric organ discharges. The premotor linkages between their electrosensory system and their medullary pacemaker nucleus, which controls the repetition rate of their electric organ discharges, appear to function differently, however. In the context of the jamming avoidance response, Eigenmannia can raise or lower its electric organ discharge frequency from its resting level. A normally quiescent input from the diencephalic prepacemaker nucleus can be recruited to raise the electric organ discharge frequency above the resting level. Another normally active input, from the sublemniscal prepacemaker nucleus, can be inhibited to lower the electric organ discharge frequency below the resting level (Metzner 1993). In contrast, during a jamming avoidance response, Apteronotus cannot lower its electric organ discharge frequency below the resting level. The sublemniscal prepacemaker is normally completely inhibited and release of this inhibition allows the electric organ discharge frequency to rise during the jamming avoidance response. Further inhibition of this nucleus cannot lower the electric organ discharge frequency below the resting level. Lesions of the diencephalic prepacemaker do not affect performance of the jamming avoidance response. Thus, in Apteronotus, the sublemniscal prepacemaker alone controls the change of the electric organ discharge frequency during the jamming avoidance response.  相似文献   

15.
Temperature manipulation has been shown to significantly affect recovery after spinal cord injury in various mammalian model systems. Little has been known thus far about the impact of temperature on structural and functional recovery after central nervous system lesions in regeneration-competent, poikilotherm organisms. In the present study, we addressed this aspect using an established model of adult spinal cord regeneration, the weakly electric teleost fish Apteronotus leptorhynchus. We observed an overall beneficial effect of increased temperature on both structural and behavioral recovery after amputation of the caudal spinal cord. Fish kept at 30°C recovered the amplitude of the electric organ discharge at more than twice the rate observed in fish kept at 22°C, within the first 20 days post-injury. This improved recovery was supported by increased cell proliferation and decreased apoptosis levels in fish kept at 30°C. The high temperature appeared to have a direct inhibitory effect on apoptosis and to lead to a compression of the duration of the wave of post-lesion apoptosis. The latter effect was presumably induced through the acceleration of the metabolic rate, a phenomenon also supported by the observation that re-growth of the tail was significantly increased in fish kept at 30°C.  相似文献   

16.
Oxygen consumption in weakly electric Neotropical fishes   总被引:2,自引:0,他引:2  
Weakly electric gymnotiform fishes with wave-type electric organ discharge (EOD) are less hypoxia-tolerant and are less likely to be found in hypoxic habitats than weakly electric gymnotiforms with pulse-type EOD, suggesting that differences in metabolism resulting from EOD type affects habitat choice. Although gymnotiform fishes are common in most Neotropical freshwaters and represent the dominant vertebrates in some habitats, the metabolic rates of these unique fishes have never been determined. In this study, O2 consumption rates during EOD generation are reported for 34 gymnotiforms representing 23 species, all five families and 17 (59%) of the 28 genera. Over the size range sampled (0.4 g to 125 g), O2 consumption of gymnotiform fishes was dependent on body mass, as expected, fitting a power function with a scaling exponent of 0.74, but the O2 consumption rate was generally about 50% of that expected by extrapolation of temperate teleost metabolic rates to a similar ambient temperature (26°C). O2 consumption rate was not dependent on EOD type, but maintenance of scan swimming (continuous forwards and backwards swimming), which is characteristic only of gymnotiforms with wave-type EODs, increased O2 consumption 2.83±0.49-fold (mean±SD). This suggests that the increased metabolic cost of scan swimming could restrict gymnotiforms with wave-type EODs from hypoxic habitats.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.  相似文献   

17.
1. An in vitro preparation of the medullary pacemaker nucleus of the weakly electric fish Apteronotus leptorhynchus was studied which fires regularly and synchronously at the fish's characteristic frequency of electric organ discharge (EOD). Upon bipolar stimulation of tissue regions through which pass prepacemaker nucleus afferents to the pacemaker, a brief, transient increase in discharge frequency ensued at short-latency (Fig. 1A). 2. Intracellular recordings revealed that the acceleration was accompanied by a depolarization and decline in action potential amplitude. The magnitude of these changes was both phase- (Fig. 5) and amplitude-dependent, with the latter showing an evident threshold effect (Figs. 4 and 12). The response was reversibly blocked by high Mg2+ saline (Fig. 1B), and the magnitude of the accelerations showed marked facilitation during repeated stimulation (Fig. 6). 3. Optical and histological identification allowed characteristically different responses in the intracellular recordings to be attributed to the two cell types of the pacemaker nucleus: pacemaker and relay cells (Figs. 2 and 3). Similar responses have been observed at these respective recording locations in the intact animal during chirping (Dye and Heiligenberg 1987). 4. Simultaneous recordings of pairs of cells revealed a transient change in the phase relationship of firing during the accelerations which was most marked between relay and pacemaker cells (Fig. 7). These dual recordings also revealed that the relay cells depolarize and accelerate more than pacemaker cells (Fig. 10), suggesting that they are the principal effectors of this behavioral modulation. 5. Trains of pulses additionally elicited a long-lasting frequency elevation which occurred at a slightly higher threshold than the brief accelerations. This slow frequency change relaxed back to baseline following a biexponential time course which closely resembled that of a distinct behavior seen in intact fish, termed 'yodeling' (Dye 1987).  相似文献   

18.
Different species have developed different solutions to the problem of constructing a representation of the environment from sensory images projected onto sensory surfaces. Comprehension of how these images are formed is an essential first step in understanding the representation of external reality by a given sensory system. Modeling of the electrical sensory images of objects began with the discovery of electroreception and continues to provide general insights into the mechanisms of imaging. Progress in electric image research has made it possible to establish the physical basis of electric imaging, as well as methods to accurately predict the electric images of objects alone and as a part of a natural electric scene. In this review, we show the following. (1) The internal low resistance of the fish’s body shapes the image in two different ways: by funneling the current generated by the electric organ to the sensory surface, it increases the fields rostrally, thus enhancing the perturbation produced by nearby objects; and by increasing the projected image. (2) The electric fish’s self-generated currents are modified by capacitive objects in a distinctive manner. These modulations can be detected by different receptor types, yielding the possibility of “electric color.” (3) The effects of different objects in a scene interact with each other, generating an image that is different from the simple addition of the images of individual objects, thus causing strong contextual effects.  相似文献   

19.
In electric fish, Apteronotus leptorhynchus, both long-term social interaction and cortisol treatment potentiates chirping, an electrocommunication behavior that functions in aggression. Chirping is controlled by the diencephalic prepacemaker nucleus (PPn-C) located just lateral to the ventricle. Cells born in adult proliferative zones such as the periventricular zone (PVZ) can migrate along radial glial fibers to other brain regions, including the PPn-C. We examined whether social interactions or cortisol treatment influenced cell addition and radial glia fiber formation by (1) pairing fish (4 or 7 days) or (2) implanting fish with cortisol (7 or 14 days). Adult fish were injected with bromodeoxyuridine 3 days before sacrifice to mark cells that were recently added. Other fish were sacrificed after 1 or 7 days of treatment to examine vimentin immunoreactivity (IR), a measure of radial glial fiber density. Paired fish had more cell addition than isolated fish at 7 days, coinciding temporally with the onset of socially induced increase in chirping behavior. Paired fish also had higher vimentin IR at 1 and 7 days. For both cell addition and vimentin IR, the effect was regionally specific, increasing in the PVZ adjacent to the PPn-C, but not in surrounding regions. Cortisol increased cell addition at 7 days, correlating with the onset of cortisol-induced changes in chirping, and in a regionally specific manner. Cortisol for 14 days increased cell addition, and cortisol for 7 days increased vimentin IR but in a regionally non-specific manner. The correlation between treatment-induced changes in chirping and regionally specific increases in cell addition, and radial glial fiber formation suggests a causal relationship between such behavioral and brain plasticity in adults, but this hypothesis will require further testing.  相似文献   

20.
A histochemical analysis of reaction rates of a series of enzymes was performed in electromotor neurons of the weakly electric fish Apteronotus leptorhynchus. These neurons were selected because of their functional homogeneity. The high metabolic activity of these cells as well as their large size facilitate cytophotometric analysis in cryostat sections. Sections were incubated for the activity of hexokinase, glucose-6-phosphate dehydrogenase, succinate dehydrogenase, NADPH dehydrogenase, NADPH ferrihaemoprotein reductase and beta-hydroxybutyrate dehydrogenase. All media contained polyvinyl alcohol as tissue stabilizer and Nitro BT as final electron acceptor. Measurements were performed with a Vickers M85a cytophotometer. Linear relationships between the specific formation of formazan (test minus control reaction) and incubation time were obtained for all enzymes although some reactions showed an initial lag phase or an intercept with the ordinate. The relatively high activities of hexokinase, succinate dehydrogenase and the extremely low activity of hydroxybutyrate dehydrogenase indicate that energy is mainly supplied by glycolysis. Glucose-6-phosphate dehydrogenase showed a high activity whereas NADPH reductase and dehydrogenase activity were low in electromotor neurons, indicating that the NADPH generated is largely used for biosynthesis. Despite their synchronous firing pattern activity, electromotor neurons showed a considerable heterogeneity with respect to their metabolic activity.  相似文献   

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