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Department of Neuroscience, Mount Sinai School of Medicine, New York, New York
Submitted 12 December 2006; accepted in final form 12 February 2007
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ABSTRACT |
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INTRODUCTION |
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). In other experimental systems, the type of stimulus does not completely specify the type of response. The response may depend on the context in which the behavior is elicited. For instance, when the cricket is in the air, stimulation of a command neuron elicits flying, whereas activation of the same neuron elicits running while the cricket is on the ground (Nolen and Hoy 1984). Context-dependent differences in behavioral responses elicited by specific command neurons were also previously observed in the medicinal leech (Esch et al. 2002). In addition to external contextual cues, application of hormones and neuropeptide modulators can influence behavioral responses elicited by the same stimulus. This was shown in the crustacean stomatogastric nervous system (Harris-Warrick and Marder 1991) and in the stridulation responses in insects (Heinrich et al. 2001). Similar mechanisms were also shown to operate in control of the feeding network in Aplysia (Jing and Weiss 2002; Jing et al. 2003; Morgan et al. 2002; Wu et al. 2003). In addition to these mechanisms, another mode of control over feeding motor programs was recently described. Proekt et al. (2004) demonstrated that previous history of activity of the feeding network also influences the subsequent motor outputs.
The feeding network in Aplysia can be activated to produce both ingestive and egestive motor programs by two input pathways: the commandlike cerebral–buccal interneuron 2 (CBI-2) and the esophageal nerve (EN) (Chiel et al. 1988; Rosen et al. 1991). However, these inputs do not unequivocally specify the nature of the motor program. CBI-2 and EN drive the motor output toward a particular steady state: ingestive in the case of CBI-2 and egestive in the case of EN. Furthermore, when the input is switched to CBI-2 after repeated stimulation of EN, the motor output does not switch immediately but remains egestive (Proekt et al. 2004).
Here, we have investigated how these slow dynamics are implemented on the interneuronal level. We focused on neurons B20 and B40 because these neurons are preferentially active in, and contribute to, egestive and ingestive responses, respectively (Jing and Weiss 2001, 2002). We found that B20 continued to fire at high frequency and B40 continued to fire at low frequency in egestive motor programs when the input was switched from EN to CBI-2. Thus firing frequency of B20 and B40 expressed the motor output, regardless of the input. Activity of another interneuron, B65, however, expressed the input regardless of the output; B65 was strongly active in EN-elicited egestive programs, but did not fire at all in egestive programs elicited immediately afterward by CBI-2. We also investigated the nature of the interaction between input- and output-representing neurons. We found that B65 produced changes in the excitability and synaptic connections of B20 and B40 and that these changes outlasted B65 firing. B65-elicited changes in B20 and B40 were similar to those that followed EN-elicited programs. These persistent changes in B20 and B40 may contribute to the persistence of the egestive responses after the input is switched from EN to CBI-2.
Taken together, our results suggest that the population of interneurons within the behavior-generating network separately encodes the higher-order input used to activate it, as well as the behavioral response elicited by that input. Furthermore, the slow dynamics of the feeding network may at least in part be reflected in the slow modulatory actions that input-representing neurons exert on output-representing neurons and their synapses.
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METHODS |
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Experiments were performed on Aplysia californica obtained from either Marinus (Long Beach, CA) or the University of Miami Aplysia Resource Facility (Miami, FL). Aplysia were maintained in circulating artificial sea water (ASW) made from Instant Ocean (Aquarium Systems, Mentor, OH), at 14–15°C. Animals weighing 150–250 g were anesthetized by injection of isotonic MgCl2 (337 mM). Buccal and cerebral ganglia were dissected out of the animal and desheathed in a dissection chamber coated with Sylgard. The ganglia were then transferred to a recording chamber, lined with Sylgard, containing about 1.5 ml of ASW (in mM: 460 NaCl, 10 KCl, 55 MgCl2, 11 CaCl2, and 10 HEPES buffer; pH 7.6). During all experiments, buccal and cerebral ganglia were maintained at 14–17°C and continuously perfused with ASW at the rate of about 0.3 ml/min. Polysynaptic activity was reduced using a high concentration of divalent cations (HiDi) solution (in mM: 420 NaCl, 10 KCl, 121 MgCl2, 13.8 CaCl2, and 10 HEPES buffer; pH 7.6) (Trudeau and Castellucci 1992).
Standard intracellular and extracellular recording techniques were used. Intracellular electrodes were filled with an electrolyte containing 2 M K acetate and 100 mM KCl and beveled to the resistance of 6–10 M
. Intracellular recordings were performed using either the Axoclamp 2B amplifier (Molecular Devices, Foster City, CA) or Getting 5A amplifiers (Getting Instruments, Iowa City, IA). All neurons were identified as described previously (Jing and Weiss 2001, 2002; Kabotyanski et al. 1998). Extracellular recordings were performed by applying suction electrodes constructed from polyethylene tubing on buccal nerves. Signals were then amplified by an AC amplifier model 1700 (A-M Systems, Carlsborg, WA).
All recordings were acquired using a Digidata 1322A (Molecular Devices) data acquisition system at the sampling rate of 5 kHz and recorded on a PC computer (Dell, Houston, TX). Digitized recordings were then plotted using Sigma Plot 5. Statistics were performed using either SigmaPlot 5 (SPSS) or Excel (Microsoft) software.
Statistical analysis
Unless otherwise indicated, one-way ANOVA was used to establish statistical significance. ANOVA was performed using Prism software package (GraphPad Software, San Diego, CA). The ANOVA was followed by a post hoc two-tail t-test with Bonferroni correction. The data sets directly compared in the post hoc analysis are indicated by brackets in the corresponding figures (*P < 0.05, **P < 0.01, ***P < 0.001).
Characterization of feeding motor programs in Aplysia
During the consummatory phase of feeding, Aplysia produces stereotyped behaviors that were earlier described as either ingestive or egestive (Kupfermann 1974a,b). In both behaviors Aplysia first protract and then retract their radula, a food-grasping organ. During ingestive behaviors, the radula closes in the retraction phase to pull food in, whereas during egestive behaviors, the radula closes in the protraction phase to push materials out. Ingestive and egestive behaviors can be distinguished using extracellular recordings obtained in vivo by differences in coordination between the activity of neuron B8 and the activity of neurons that produce radula retraction (Morton and Chiel 1993a,b). The central pattern generator (CPG) that produces these feeding behaviors is located in the buccal ganglion. In vitro this feeding CPG is capable of producing coordinated patterns of activity referred to as feeding motor programs. Although not identical, these in vitro elicited motor programs share essential characteristics with patterns of activity recorded in vivo. During ingestive motor programs, the radula closer motoneuron B8 is predominantly active in the retraction phase, whereas during egestive motor programs, B8 is predominantly active in the protraction phase (Jing and Weiss 2001, 2002; Morgan et al. 2002; Proekt et al. 2004). Based on previous findings, we defined the protraction phase as activity recorded in the I2 nerve (Hurwitz et al. 1996; Jing and Weiss 2001 2002; Morgan et al. 2002; Nargeot et al. 1999a,b). The retraction phase was defined as high frequency of activity recorded in the buccal nerve 2 (B.n.2) after the end of activity recorded in the I2 nerve (Morton and Chiel 1993a,b; Nargeot et al. 1999a,b).
The differences in the coordination between the radula protraction, retraction, and closure activity in ingestive and egestive motor programs are clearly seen when the firing frequency of B8 during the protraction phase is plotted against the firing frequency of B8 in the retraction phase. Morgan et al. (2002) demonstrated that, in this plane, ingestive and egestive motor programs form two clusters (Fig. 1). Approximate cluster boundaries are shown as gray ovals; representative recordings of ingestive and egestive motor programs are shown in their corresponding clusters.
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), which is located in the cerebral ganglion, or the EN (Chiel et al. 1988; Morton et al. 1991) that connects the esophagus and the gut with the buccal ganglion. The CPG for feeding-related responses is located in the buccal ganglia and, except for CBI-2, all of the neurons studied herein are located in the buccal ganglion. Both CBI-2 and EN are thought to be important in feeding behavior and are capable of eliciting motor programs (Due et al. 2004; Evans et al. 2003; Horn et al. 2004; Jing and Weiss 2005, 2002, 2005; Morgan et al. 2002; Proekt et al. 2004; Rosen et al. 1991) and functional ingestive and egestive responses (Chiel et al. 1988; Church and Lloyd 1994; Jing and Weiss 2005; Zhurov et al. 2005) in semi-intact preparations. CBI-2 is known to be activated by touching food (e.g., seaweed) to the lips of the animal (Jing and Weiss 2005; Morgan et al. 2002; Rosen et al. 1991), whereas the EN conveys sensory information concerning the stretch of the esophagus (Kuslansky et al. 1987; Morgan et al. 2002) (although it may also convey other signals). CBI-2 was stimulated to fire action potentials at 9 Hz for the duration of the protraction phase. Short (20- to 30-ms duration), square current pulses, each of which triggered a single action potential, were used. No DC predepolarization was used during CBI-2 stimulation. Because DC prestimulation is required to recruit CBI-3 activity (Morgan et al. 2002), very little if any CBI-3 firing was recruited during CBI-2–elicited motor programs. EN was stimulated at about 2 Hz using 3-ms current pulses delivered by an extracellular electrode. All stimulations were performed using a Grass S48 Stimulator (Grass Instruments, Quincy, MA). |
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RESULTS |
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). In the following experiments, using the experimental paradigm developed by Proekt et al. (2004), we characterized how this history dependency is reflected in the activity of several CPG interneurons.
We first focused on interneurons B20 and B40 because activity of these neurons is critical for the expression of egestive and ingestive motor programs, respectively (Jing and Weiss 2001, 2002). We repeatedly stimulated CBI-2 throughout the protraction phase with an interprogram interval of 30 s. Nine motor programs were elicited in this fashion. As shown previously (Proekt et al. 2004), after nine consecutive CBI-2 stimulations at 30-s intervals the system settles into a steady state defined by consistent ingestive responses elicited by CBI-2 stimulation. A representative example of one such ingestive response is shown in Fig. 2A : CBI-2 control. The EN was then stimulated for 5 min; this stimulation routinely elicited nearly 10 motor programs. In these egestive motor programs, B8 fired predominantly in the protraction phase and B20 fired at higher frequency than in ingestive CBI-2–elicited motor programs (Fig. 2A, box). We then switched the stimulation back to CBI-2. Because of the slow dynamics, the network disregarded the switch in the stimulus and continued to generate egestive motor programs (Fig. 2A, CBI-2, 0 min). The firing frequency of B20 also did not switch immediately; B20 continued to fire at higher frequency than it did in CBI-2–elicited ingestive programs. On repeated CBI-2 stimulation, the motor programs once again became ingestive (Fig. 2A, CBI-2, 5 min). This was accompanied by a decrease in the firing frequency of B20 back to control levels.
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Grouped data for the two preceding experiments (Fig. 3A) demonstrate that changes in B20 (filled circle) and B40 (empty circle) firing frequencies paralleled changes in the ingestive versus egestive characteristics of the motor output, as measured by B8 firing frequency in the protraction (filled square) and the retraction (empty square) phases (Fig. 3B). Specifically, the increase in the B20 firing frequency paralleled the increase in the B8 firing frequency in the protraction phase, whereas the decrease in the B40 firing frequency paralleled the decrease in the B8 firing frequency in the retraction phase. In Fig. 3C, we replotted the same data as in Fig. 3B to demonstrate that these changes in the firing frequency of B8 amount to qualitative changes in the nature of the motor programs. The history-dependent changes in B8 firing shown in Fig. 3, B and C were similar to those shown in Proekt et al. (2004). Taken together, the data shown in Fig. 3 indicate that the firing frequency of B20 and B40 tracked the slow dynamics of the motor output of the feeding network.
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; Jing and Weiss 2005; Kabotyanski et al. 1998), showed a dramatically different pattern of firing in the same experimental paradigm. In ingestive motor programs elicited by CBI-2 before EN stimulation, B65 fired very weakly in the protraction phase (Fig. 4A1), whereas in EN-elicited egestive motor programs B65 fired robustly during the protraction phase (Fig. 4A2). When the stimulus was switched back to CBI-2, although the network continued to generate egestive motor programs, B65 firing was immediately shut down (Fig. 4A3). Grouped data for this experiment (Fig. 4B) show that B65 fired at the same frequency in motor programs elicited by CBI-2, irrespective of whether they were ingestive or egestive. In contrast, B65 fired at very different frequencies in similar egestive motor programs elicited by EN and CBI-2. Thus in contrast to B20 and B40, the firing frequency of B65 tracked the input that activates the CPG, regardless of the motor output elicited by this input.
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) and one of the defining features of egestive motor programs is that B8 fires predominantly during the protraction phase (Morgan et al. 2002). B20 is known to contribute to this egestive firing pattern of B8 by exciting it during the protraction phase by a direct synaptic connection (Diaz-Rios and Miller 2005; Jing and Weiss 2001; Proekt et al. 2004). Figure 5A demonstrates that, as previously shown by Proekt et al. (2004), EN stimulation also leads to the persistent potentiation of the B20-to-B8 synapse. The increase in the excitability of B20 and the potentiation of the B20-to-B8 synapse may act synergistically to increase the excitatory drive to B8 during the protraction phase and thus may contribute to the persistence of egestive motor programs after the switch in the stimulus.
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). In ingestive motor programs B8 fires predominantly in the retraction phase (Morgan et al. 2002). B40 promotes this ingestive firing pattern of B8 by producing a complex postsynaptic potential (PSP) in B8 that consists of the fast inhibitory postsynaptic potential (IPSP) and a slow excitatory postsynaptic potential (EPSP) that outlasts B40 firing (Jing and Weiss 2002; Jing et al. 2003). Whereas the fast IPSP suppresses B8 firing in protraction, the slow EPSP promotes B8 firing in the retraction phase (Jing and Weiss 2002; Jing et al. 2003). In the experiment shown in Fig. 6C, we stimulated B40 to fire a fixed number of action potentials and characterized the resulting EPSP before and after EN stimulation. Because the slow EPSP tends to elicit action potentials in B8, B8 was hyperpolarized to about –80 mV (Jing et al. 2003) during this experiment; thus only the excitatory component of the B40-to-B8 PSP is seen in these recordings. After EN stimulation, the slow EPSP was suppressed. This suppression recovered over the course of 3 min (Fig. 6, C and D). Thus repeated stimulation of EN elicits persistent potentiation of neuronal mechanisms that promote egestive motor programs and coordinated persistent suppression of mechanisms that promote ingestive motor programs.
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). We injected current pulses into B40 as in Fig. 6A. After several control current injections, during the 30-s interval between these current injections, we stimulated B65 to fire action potentials at 10 Hz for 20 s. Three such bursts of action potentials were elicited in B65. B65 stimulation resulted in a decrease in the excitability of B40 (Fig. 7A, representative recordings; Fig. 7B, grouped data). This decrease in the excitability of B40 recovered over the course of 90 s and appeared to be elicited by B65 by a monosynaptic pathway because the stimulation of B65 in solution containing HiDi also resulted in a decrease in the excitability of B40 (n = 3; data not shown).
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We then characterized the effect of B65 stimulation on the excitability of B20 in a similar paradigm and found that this resulted in the progressive increase in B20 excitability (Fig. 8A, representative recordings; Fig. 8B, grouped data). This increase in the excitability dissipated within 30 s after the end of B65 stimulation.
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DISCUSSION |
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suggested that state dependency may be mediated by the plasticity of the synaptic connections of the output-representing neuron B20. Here, we have extended these observations to show that plasticity associated with state dependency also involves other neurons and synaptic connections. Our results further suggest that these coordinated changes in multiple loci within the feeding network may, in part, be orchestrated by the input-representing neuron B65.
Previous work identified many neurons within the feeding circuitry of Aplysia (reviewed in Cropper et al. 2004). Firing of some neurons is important for the initiation and maintenance of the protraction and the retraction phases, irrespective of whether the motor program is ingestive or egestive (Hurwitz and Susswein 1996; Hurwitz et al. 1996, 1997; Jing and Weiss 2001). These neurons were not directly investigated here and are shown schematically as mutually inhibitory P and R neurons (Fig. 9B). Yet, other neurons shape the egestive versus ingestive nature of the motor programs. Several lines of evidence point to the importance of B20 and B40 in shaping these ingestive versus egestive characteristics: 1) B20 is more active in egestive motor programs, whereas B40 is more active in ingestive motor programs; 2) stimulation of B20 leads to the generation of more egestive motor programs, whereas stimulation of B40 results in more ingestive motor programs; and 3) suppression of B20 disrupts the generation of egestive motor programs, whereas suppression of B40 disrupts ingestive motor programs (Jing and Weiss 2001, 2002). Although P neurons directly excite B20 and B40 by electrical and chemical synapses (Fig. 9B) (Jing and Weiss 2001, 2002), they are not thought to contribute to specifying the ingestive versus egestive nature of the motor programs because they fire similarly during ingestive and egestive programs (Jing and Weiss 2001, 2002),
We find that firing of both B20 and B40 was persistently altered after repeated EN stimulation (compare Fig. 9, B2 and B3). In EN-elicited egestive programs, B20 fired at high frequency (Fig. 2A), whereas B40 fired at low frequency (Fig. 2B). After the input was switched from EN to CBI-2, B20 firing frequency remained high, whereas B40 firing frequency remained low. Consistent with the persistent changes in the firing frequency of B20 and B40, we found that the excitability of these neurons was persistently altered after repeated EN stimulation—B20 excitability was increased (Fig. 5), whereas B40 excitability was decreased (Fig. 6). Thus the persistence of egestive responses after the switch in the stimulus is accompanied by both the potentiation of processes that promote egestion and the concurrent suppression of processes that promote ingestion.
This conclusion is further supported by experiments with the synapses through which B20 and B40 act to shape the ingestive versus egestive characteristics of the motor output. We previously demonstrated that the B20-to-B8 synapse is facilitated after repeated EN stimulation and that this synaptic facilitation is accompanied by an increase in the functional efficacy of the synapse (Proekt et al. 2004) (Fig. 9, B2 and B3). This synapse serves to excite B8 during the protraction phase. Because high frequency of B8 firing in the protraction phase is a defining characteristic of egestive responses (Morgan et al. 2002), the potentiation of the B20-to-B8 synapse serves to promote egestion.
Here we have shown that the excitatory component of the B40-to-B8 synapse was suppressed after EN stimulation. B40 fires during the protraction phase of ingestive programs, but promotes B8 firing in the subsequent retraction phase (Fig. 9B1). B40 accomplishes this by eliciting a complex synaptic response in B8 that consists of the fast inhibitory component and the slow excitatory component. Although the fast inhibition serves to suppress B8 firing during the protraction phase, the slow excitation outlasts B40 firing and promotes B8 firing in the retraction phase (Jing and Weiss 2002; Jing et al. 2003). The high firing frequency of B8 in the retraction phase defines ingestive responses (Morgan et al. 2002). Thus suppression of the excitatory component of the B40-to-B8 synapse serves to suppress ingestive responses after repeated EN stimulation.
These experiments illustrate that the reconfiguration of the feeding network by repeated EN stimulation involves simultaneous coordinated changes in several neurons and synapses. Similar conclusions have been reached in the analysis of network reconfiguration in the leech CPG that produces crawling and swimming motor programs. By simultaneously observing the firing of a population of neurons, Briggman et al. (2005) were able to effectively predict which motor pattern would be produced, earlier than by observing any individual neuron. Here, we demonstrate that these coherent changes in firing of a population of neurons are paralleled by changes in their excitability and synaptic connections.
Our experiments further suggest that firing of neuron B65 can produce these coordinated changes in neuronal excitability and synaptic connections. Stimulation of B65 resulted in the persistent increase in the excitability of B20 (Fig. 9B2). Potentiation of the B20-to-B8 synapse is known to be activity dependent (Proekt et al. 2004); thus the increase in the excitability and the consequent increase in the firing frequency of B20 may also indirectly lead to the potentiation of the B20-to-B8 synapse (Fig. 9B2). Additionally, B65 firing leads to the persistent suppression of the B40 excitability and the direct suppression of the B40-to-B8 synapse.
B65 synapses onto many CPG neurons and receives input from them (Due et al. 2004; Jing and Weiss 2005; Kabotyanski et al. 1998). Stimulation of B65 can elicit motor programs (Kabotyanski et al. 1998). Furthermore, suppression of B65 firing produces a modest effect on the protraction duration and B8 firing frequency in EN-elicited egestive motor programs (Due et al. 2004). Thus B65 is a component of the CPG. Here we found that B65 did not fire during ingestive programs. Furthermore, although B65 was strongly active in EN-elicited egestive programs, it did not fire in egestive programs elicited by CBI-2. Thus B65 firing is not essential for egestive programs—the only type of motor output that recruits its activity. The lack of firing of B65 in CBI-2–elicited motor programs may be explained by direct inhibition of B65 by CBI-2 (Jing and Weiss 2005).
Thus although B65 is a bona fide component of the CPG, its firing does not have strong significance for the immediate motor output. Instead, activity of B65 most closely followed the input. It remains to be determined, however, whether additional neurons are also selectively activated in EN-elicited programs. This is an interesting question because the EN is the only nerve that connects the feeding network with the gut and the esophagus. As such, EN is likely to carry information about the presence of chemo-stimuli in the gut (Susswein et al. 1984) and about the degree of stretch of the gut and the esophagus (Kuslansky et al. 1987; Morgan et al. 2002). It is possible that these different sensory modalities recruit activity of distinct populations of CPG elements. In addition, EN is known to contain neuropeptides and two of these neuropeptides have been shown to convert ingestive CBI-2–elicited motor programs to egestive ones (Jing et al. 2007; Wu et al. 2006). If these neuropeptides are released during EN stimulation, they could exert persistent action on CPG elements. Through this mechanism, neuropeptides released by EN stimulation could provide a parallel means of input representation in the feeding CPG.
Previous work that examined how the same CPG produces different behaviors focused on identifying command neurons that activate the CPG (e.g., Beenhakker and Nusbaum 2004 Croll et al. 1985; Morgan et al. 2002; Rosen et al. 1991). Previously it was argued that control over which behavior is produced is achieved by a population code (Georgopoulos et al. 1986; Kristan and Shaw 1997; Lewis 1999; Lewis and Kristan 1998), meaning that a population of higher-order neurons activated by the stimulus determines the nature of the response. A similar scheme was proposed for Aplysia feeding (Jing and Weiss 2005; Morgan et al. 2002). Nonetheless, in the feeding system, the decisions as to which behavior to generate is controlled not just by the population of higher-order neurons, but also by the state of the feeding network itself. By focusing on this state dependency, we discovered two fundamentally distinct populations of neurons—input- and output-representing interneurons. This distinction is well established in networks involved in decision making. For instance, when a monkey is trained to indicate the direction of movement of dots on a screen by a saccade, the activity of neurons in the extrastriate visual cortex reflects the net movement of dots, whereas the activity of neurons in the frontal eye fields reflects the direction of the saccade (Cohen and Newsome 2004). Interestingly, electrical stimulation of the frontal eye fields immediately before the voluntary saccade elicits a saccade that deviates from the control stimulation in the same direction as the subsequent voluntary action (Gold and Shadlen 2000). This suggests that the behavior-generating neurons are reconfigured before the onset of action and this reconfiguration dictates the nature of the subsequent behavioral response.
There is considerable debate concerning the role of individual brain areas in decision making. For instance, it was argued that neurons in the lateral intraparietal area (LIP) encode the perception of motion (Assad and Maunsell 1995; Williams et al. 2003), visual attention (Bisley and Goldberg 2003), decisional processes (Shadlen and Newsome 2001), and forming intentions (i.e., plans for future behavior) (Andersen and Buneo 2002). Here, we used the simplicity of the invertebrate preparation to show that firing of B65 reflects the instantaneous stimulus, although its actions may shape the future behavior of the feeding network. Thus firing of B65 may provide a link between the instantaneous stimulus and future behaviors.
Virtually all animals, from simple invertebrates to humans, live in complex environments. Animals' survival depends on their ability to respond differentially and efficiently to the different stimuli to which they are exposed. However, making an animal respond immediately to a stimulus may leave it at the mercy of transient stimuli and prevent it from achieving a specific goal. This may have led to evolution of strategies that allow animals to balance the need to respond to sensory cues with the need to pursue their internal goals. The slow dynamics of switching between different behavioral responses could serve to suppress responses to transient stimuli and emphasize responses to persistent and repeated stimuli. Because environmental stimuli are not distributed randomly, but are clustered or clumped together, the slow dynamics exhibited by the feeding CPG could embody an adaptive behavioral strategy.
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GRANTS |
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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Address for reprint requests and other correspondence: K. R. Weiss, Department of Neuroscience, Mount Sinai School of Medicine, One Gustave L. Levy Place, Box 1065, New York, NY 10029 (E-mail: Klaudiusz.Weiss{at}mssm.edu)
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