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First published online August 9, 2007
Journal of Experimental Biology 210, 2836-2842 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.003988
Male discrimination of receptive and unreceptive female calls by temporal features
1 Neurobiology and Behavior, Columbia University, New York, NY,
USA
2 Biological Sciences, Columbia University, New York, NY, USA
* Author for correspondence (e-mail: taffeta{at}berkeley.edu)
Accepted 10 June 2007
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Summary |
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Key words: anura, auditory discrimination, categorical perception, frog, vocal communication, Xenopus
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Introduction |
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TOPSummaryIntroductionMaterials and methodsResultsDiscussionReferences |
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). In
categorical perception – used, for example, in the Polynesian field
cricket's identification of conspecific mating calls and predator bat
echolocation calls – differences along a continuum are perceived with
greater acuity at category boundaries than within the range of one category
(Wyttenbach et al., 1996).
Categorical perception can improve the speed and accuracy of discrimination
between different classes of social communication signals
(Gleason and Ratner, 1998).
However, where there is an overlap between signals whose significance differs,
a continuous strategy may be more appropriate.
The ticking and rapping calls (Tobias
et al., 1998) of the South African clawed frog, Xenopus
laevis (Daudin), provide an informative system in which to investigate
the relative roles of continuous and categorical strategies in acoustic
communication. X. laevis communicates underwater at night using eight
vocal call types made up of clicks that differ in temporal patterns
(Tobias et al., 2004). The two
female calls are rapping, a rapid trill, which functions as an acoustic
stimulant that increases male advertisement calling, and ticking, a slower
trill, which suppresses male vocal production
(Tobias et al., 1998).
Sexually receptive females rap whereas unreceptive females tick.
Other X. laevis calls differ in click envelope, in relative
amplitude between clicks, and/or in trill durations, but the most distinctive
acoustic difference between female rapping and ticking is click rate
(Tobias et al., 1998). The
mean interclick interval (ICI) of rapping is 81 (±14 ms s.d.) and the
mean of ticking is 219 (±71 ms), with some overlap in range (between 50
and 175 ms, Fig. 1A)
(Tobias et al., 1998).
Regardless of the female call type or the individual, spectral energy in the
broadband clicks peaks at 1.2 kHz (Tobias
et al., 1998). Rise-times and fall-times appear only slightly
sharper in ticking clicks than in rapping clicks. A solitary female often raps
at levels similar to ticking, 39 dB with reference to 1 µPa, but during a
male–female duet in our 2 m laboratory tank the maximum rapping sound
level reached 86 dB re 1 µPa (Tobias et
al., 1998). Males use click rate to distinguish the sex of callers
(Tobias and Kelley, 1987;
Vignal and Kelley, 2007).
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Materials and methods |
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). Labeling boundary playback experiments were performed in a
fiberglass tank (1 mx2 mx0.5 m deep) at the beginning of the dark
period, under dim indirect illumination from a red lamp. Dishabituation
experiments were performed in an approximately 1 m3 fiberglass tank
filled 3/4-full with dechlorinated, filtered water and resting on a layer of
foam. Labeling boundary experiments took place during the springs of 2003 and
2004, and dishabituation experiments during the fall and winter of 2005-2006,
at water temperatures from 19 to 23°C.
Stimuli
In the labeling boundary experiment, stimulus CDs were played on a Sony
Walkman (D-EJ360; Tokyo, Japan) or a CD recorder (Marantz CDR300; Mahwah, NJ,
USA). In the dishabituation experiment, the initiation and insertion of sound
stimuli was instead controlled on-line on a laptop computer (Apple Titanium
G2). Call stimuli were amplified (Realistic MPA30) and presented through an
underwater loudspeaker (University Sound UW-30 Diatran underwater loudspeaker,
frequency response 0.1–10 kHz; San Diego, CA, USA). The volume of
playback was matched to recordings of a live female ticking in the same tank
during pilot experiments. To create stimuli, we started with bouts of female
ticking (selection in Fig. 1B)
recorded in the field (Tobias et al.,
1998), using a Cornell Bioacoustics Program hydrophone (output
sensitivity –163±3 dB re 1 V/µPa, frequency sensitivity
0.015–10 kHz; Ithaca, NY, USA), into a Marantz cassette tape recorder
(PMD430) and digitized. We edited periods of silence between the clicks to
produce the following constant intervals: 81, 98, 120, 160, 180 and 219 ms
ICIs (see Fig. 1A,C).
Playback procedures
Labeling boundary protocol
The click rates presented include the means for rapping (81 ms ICI) and for
ticking (219 ms ICI). In addition, we presented three intermediate rates
– 98, 120 and 160 ms ICIs – that represent log intervals between
the means of rapping and ticking (Fig.
1A) (Tobias et al.,
1998). Logarithmic spacing of the intermediate stimuli was chosen
because human listeners and European starlings perceive rate logarithmically,
with similar behavioral discrimination over logarithmically proportional
intervals (Braaten and Hulse,
1993; Palmer and Krumhansl,
1990).
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Every subject was tested with all three intermediate stimuli. Each intermediate stimulus was presented nightly for a single block of 5 nights. The order of the three testing blocks was determined pseudorandomly. For most blocks, we analyzed data from nights 1–4. In some blocks, subjects did not call robustly during initial stimuli on the first night, possibly because of novelty-induced suppression. If time spent calling to the control stimuli (rapping and ticking click rates) on night 1 totaled <30 s, we analyzed nights 2–5. This occurred in 16 out of 48 blocks, involving nine subjects. Responses were averaged over the 4 nights. To confirm that nights 2–5 in the 16 blocks were equivalent to nights 1–4 in the other blocks, we performed a repeated-measures analysis of variance (RM-ANOVA) testing the total calling (responses and baseline) by night. Nights 4 and 5 each produced significantly more calling than night 1 (P<0.05 and P<0.01, respectively). No significant difference between nights 1 and 2 was found. A week without hormone injections preceded each of the three testing blocks.
Dishabituation protocol
The phenomenon of dishabituation indicates discrimination between stimuli
that lie nearby along the gradient of a perceptual parameter. We used a
dishabituation paradigm (Fig.
3A) to test for discrimination between two click rates labeled as
ticking (180 ms and 219 ms ICI). These two rates were chosen because 219 ms
ICI is the mean and therefore likely to be an effective suppressor; 180 ms ICI
is more similar to rapping but is well within the range labeled as ticking.
After suppression, when frogs had habituated to one or the other of the
ticking stimuli by beginning to call again with three fast trills of the
advertisement call (Fig. 3B),
we either switched the stimuli (test trial,
Fig. 3A) or held the stimulus
constant throughout the habituated calling (control trial). In either
condition, time spent calling was counted until a pause of at least 30 s
occurred. If the frog changed his calling behavior in any way – whether
by suppressing more quickly, or by calling longer or with more rapid trill
alternations – this change would constitute dishabituation and would
indicate that the frog had detected the difference in stimuli and was altering
his calling accordingly.
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Experiments were begun after the male started calling (either spontaneously or, more often, promoted by 30 s of clicks broadcast at the rapping rate after 15 min without calling). After the male had produced three fast trills of the advertisement call, we presented either the 180 or the 219 ms ICI stimulus chosen by coin flip (Fig. 3A). We measured time spent calling (all male calls) until there was a 30 s pause in calling, which was our criterion for suppression. Time from the stimulus onset until the beginning of the 30 s suppression period is the time to suppression (TTS; Fig. 3A).
After two trials measuring the TTS to the two stimuli, we began dishabituation trials (Fig. 3). These trials began in the same way as the introductory trials, with the exception that we continued to play the stimulus past the 30 s period of suppression. When a male again made three fast advertisement trills, we considered this calling as an instance of habituation. We calculated the time to habituation (TTH) as the time following TTS up until the male started calling again (Fig. 3A). (Habituation is defined as the decreased responsiveness to repeated stimulus exposure; in this case, the response that habituates is the suppression response, rather than the vocalization itself.) The TTS and TTH criteria were originally defined based on suppression and bout times in pilot experiments with three males (data not shown). In test trials (Fig. 3), after the three fast trills of habituated advertisement calling, the stimulus then playing was switched to the other stimulus (180 or 219 ms ICI) using the stimulus-controlling computer. A custom-made stimulus program used the audio environment Max/MSP 4.5 (Cycling'74, San Francisco, CA, USA) to make a patch that looped broadcasts of habituation stimuli and switched smoothly between stimuli during dishabituation trials. In control trials, there was no stimulus change; the stimulus continued until the male stopped calling for 30 s. Measurements of the male's response to test and control trials included the time spent calling, the total time until a 30 s period without calling, and the number and rate of fast advertisement trills. If the male responded differently to test trials than to control trials, this alteration in vocal behavior would constitute dishabituation and would indicate that the male had detected the change in click rate.
Conditions were presented in a pseudorandom order, balanced to include all four combinations that differed in which stimulus came first and whether the stimulus changed mid-bout upon habituation. Trials were aborted after 10 min of suppression. Males were tested on successive nights (1 to 3) until two sets of four-stimulus combinations had been tested; 19 males completed the stimulus set.
Data collection and analysis
Male vocal responses were recorded with two hydrophones (Cornell
Bioacoustics Program, sensitivity as described above), one suspended in the
center of the tank at a depth of 0.5 m and the other placed on the bottom of
the tank slightly to one side of the speaker in the corner. The hydrophone
channels were recorded to CD (Marantz CD recorder CDR300) or stored as stereo
MP3 files on a digital recorder (Marantz PMD670).
Male vocalization recordings were analyzed using Goldwave on a PC (Hewlett Packard Pavilion). Time spent calling was determined to within 0.5 s by visual inspection of the files or, when the recording failed because of CD error, by using the clock on the CD recorder and transcribing calling times to the nearest second by ear. In the labeling boundary experiment, isolated clicks were rounded up to 0.5 s.
The male advertisement call alternates between short, fast trills that are
strongly intensity-modulated, and long, slow trills that vary less in peak
click amplitudes (Fig. 3B).
Bouts of advertisement calling sometimes include an acoustically related call,
male answer calling, that differs in the relative duration of fast and slow
trills: fast trills are lengthened and slow trills are shortened
(Tobias et al., 1998). Males
answer call during duets with rapping and ticking females, as well as during
interactions with other males (Tobias et
al., 1998; Tobias et al.,
2004). Because answer calls occur during acoustic stimulation or
physical interactions between animals, whereas advertisement calling is also
given by isolated males without stimulation, we analyzed the number of fast
trills per unit of time as a possible indication of male changes in perception
or motivation. For the dishabituation experiment, fast advertisement trills
were tallied, and inter-fast-trill intervals (IFTIs) were measured to test for
dishabituation.
To avoid observer bias in the test for dishabituation, measurement of IFTIs in the habituated bouts was assessed visually by a second observer unaware of the stimulus code. Although the stimulus changes are audibly apparent in the sound files, the much higher sound level of the male call prevented the observer from seeing stimulus changes during the visual inspection of the signal files in Goldwave (waveforms depict amplitude versus time in Fig. 3B). To determine whether a change had occurred in the rate of fast trills after the stimulus switch, the second observer compared the times between the first three habituated fast trills (two intervals) and the subsequent three fast trills (two intervals). In the case of control trials the latter two intervals occurred after the stimulus would have been switched had it been a test trial. The third fast trill interval was excluded because the stimulus switch occurred at different time points within this slow trill (Fig. 3B).
In both experiments, two-tailed RM-ANOVAs were used to test for
significance at the P<0.05 level. Fisher's probable least squares
difference (PLSD) was used as a post-hoc test because the probability of type
I error was deemed acceptable at 
=0.05. All values given are means
± s.e.m.
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Results |
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TOPSummaryIntroductionMaterials and methodsResultsDiscussionReferences |
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). Whereas
ticking suppresses a male when it is presented during a bout of advertisement
calling (see dishabituation experiment, below), it does not suppress calling
relative to the baseline condition of males alone in a tank without acoustic
playback at the start of the experiment. Of the three intermediate stimuli (Fig. 4), the one with a 98 ms ICI stimulated as much calling as the rapping stimulus (81 ms ICI), but significantly more calling than the ticking stimulus (219 ms ICI) presented on the same nights (P<0.0005, RM-ANOVA, Fisher's PLSD, Fig. 4A). The response to the 160 ms ICI stimulus was significantly lower than the response to rapping on the same nights (P<0.05, Fisher's PLSD, Fig. 4C). By contrast, calling during the 120 ms ICI stimulus did not differ significantly from calling during rapping or from calling during ticking in that block (P>0.05, RM-ANOVA, Fig. 4B). A significant effect of the call with 120 ms ICIs might have been masked by variability between individual males; for example, if half the animals responded `positively' (as to rapping) and half `negatively' (as to ticking or as in baseline calling). However, the variability in response across nights for each frog was equivalent to the variability between individuals (the by-subject analysis within the RM-ANOVA was not significant).
There was no significant stimulus order effect: the amount of time spent calling during the intermediate test stimuli (98, 120 or 160 ms ICI) did not depend on whether the 81 ms or the 219 ms ICI control stimulus preceded (t-test rejected at P>0.05, Fig. 5).
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Male suppression and habituation to female ticking
Both stimuli within the range of ticking ICIs effectively suppressed male
calling (Fig. 6, left-hand
panel), as measured by time to suppression or TTS (180 ms ICI:
91.3±72.0 s; 219 ms ICI: 80.5±101.5 s; N=20; RM-ANOVA
rejected at P>0.05).
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115 s (Fig. 6,
right-hand panel). The median TTH was not significantly different between the
two stimuli (RM-ANOVA rejected at P>0.05). An example of a male
vocalization during habituation is shown in
Fig. 3B.
Absence of dishabituation to different click rates of female ticking
There was no significant difference in total amount of time spent calling
during the habituated bouts (Fig.
3B) whether the stimulus had changed or not, nor was there a
difference in the bout lengths (all comparisons rejected at
P>0.05, RM-ANOVA). Mean habituated bout lengths were
139±167 s to 180 ms ICI, 122±105 s to 219 ms ICI, 124±97
s when the stimulus was switched from 180 to 219 ms ICI, and 165±238 s
when the stimulus was switched from 219 to 180 ms ICI. Neither was there a
significant difference in the number of fast advertisement trills, which was
similar across stimuli (median 35 trills in the control trials, and 36 trills
in the test trials), or in the number of fast trills per total habituated
calling (0.7 fast trills s–1 for both control and test
trials). Was there any more immediate change in fast trill rates just before
and after the time of stimulus change? The two IFTIs after the stimulus switch
(or after a comparable time point in control trials) tended to be longer than
the first two IFTIs in the habituated bout (2.2±0.8 versus
1.4±0.05 s IFTI), but there was no significant difference
(P>0.05, RM-ANOVA). We also measured the IFTIs for spontaneous
calling in the labeling boundary experiment. Across the first five trills in
one bout of spontaneous calling the IFTI speeds up by 46 ms (903±200 ms
mean and s.d. versus 857±161 ms; P<0.05). We would
thus expect the IFTI to be sensitive to changes that could occur during a
switch in stimuli in the dishabituation experiment. No such change was
detected.
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Discussion |
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TOPSummaryIntroductionMaterials and methodsResultsDiscussionReferences |
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). For the isolated male, both ticking and rapping
stimulate calling above baseline, but rapping is more effective
(Vignal and Kelley, 2007),
also shown in the present study. The average ICI of ticking is 219 ms but the
range of ICIs during ticking bouts is broad
(Fig. 1A). The average ICI of
rapping is 81 ms and has a more narrow range of variation than ticking. The
variability in ICIs produces ambiguous signals in the range of click rate
values that overlap between rapping and ticking. We thus sought to determine,
using the labeling boundary experiment, whether males identify intermediate
stimuli (in the overlapping range) as rapping or ticking, using the amount of
time spent calling as a measure. Given the click rates that males clearly label as ticking, do males discriminate within the ticking range? We found that the male calling response to constant ticking suppresses. The suppression response eventually habituates, which is to say that with continued presentation of a ticking stimulus, males resume calling. We compared two ICIs within the ticking range for their effectiveness as suppressors and as habituating stimuli. Then we tested for dishabituation to determine whether males responded differentially to a change in ticking ICI.
The labeling experiment thus involved click rates in common between rapping and ticking; the dishabituation experiment used click rates males labeled as distinctly ticking. We will summarize these results and compare them to what is known about categorical partitioning of continuously varying stimuli by other species.
Labeling boundary between female calls
The fastest intermediate test stimulus, 98 ms ICI, was identified by the
male as rapping, and the slowest stimulus, 160 ms ICI, was identified as
ticking. The response to 120 ms ICI did not differ from either rapping or
ticking responses. This result is not due to some males treating the 120 ms
ICI stimulus consistently as rapping whereas others responded as though to
ticking. In all other comparisons between rapping and ticking stimuli, males
respond with significantly more calling in response to rapping (see also
Tobias et al., 1998). Ticking
may have elicited more calling than usual (and rapping less) on the nights
including 120 ms ICI because prolonged exposure to an ambiguous stimulus may
have reduced the archetypal quality of ticking and rapping rates in
comparison.
Over the tested range from 81 to 219 ms ICI, there is no indication of a sharp boundary in the male responses to five click rates. Instead, the amount of time that the male spends calling appears to reflect the amount of overlap in the distribution of ICIs present in bouts of natural female rapping or ticking (Fig. 1B). Because 98 ms ICI is the upper limit of the standard deviation of natural rapping rates, whereas 160 ms is the lower limit of natural ticking, the male responds to 98 and 160 ms ICI accordingly. More intermediate rates should be ambiguous. In fact, when 120 ms ICI is played, the male's response does not differ from his response to either ticking or rapping. Relatively few ICIs from either rapping or ticking contain this interval. It is a truly ambiguous signal and the male response is correspondingly ambiguous.
Suppression response and habituation to female ticking
When a male is calling, a broadcast of ticking can produce vocal
suppression within 91 s. Within the ticking range, the two stimuli tested (180
and 219 ms ICIs) were equally effective in time to suppression; males do not
respond preferentially to these intervals.
A vocally suppressed male will resume calling if the ticking stimulus is continued (habituation). Again, the time to habituation was equivalent for the two ICIs tested; males do not respond differentially to these click rates. When the stimulus producing habituation was exchanged for the other stimulus, the male's response did not differ from the condition of no exchange. Thus, as far as can be determined from the male's vocal response, there is no differential sensitivity to ICIs characteristic of ticking even though the two ICIs tested occur occasionally also in rapping (Fig. 1A).
Continuous and categorical perception
In categorical perception, differences along a continuum are perceived with
greater acuity at category boundaries than within the range of one category.
This perceptual phenomenon requires both (1) a sharp labeling boundary between
classes and (2) a lack of discrimination between stimuli in the same class.
Results with ICIs within the ticking class (including time to suppression,
time to habituation and the absence of dishabituation) indicate that males do
not discriminate among ticking stimuli (consistent with the second requirement
for categorical perception). However, results of the labeling boundary
experiments provide no evidence of a sharp boundary between rapping and
ticking (failing the first requirement); males respond ambiguously at the
labeling boundary between the two calls, around 120 ms ICI.
If the perceived boundary between rapping and ticking were sharp (i.e. if nearby click rates on opposite sides of some boundary rate produced significantly different responses), males might distinguish female calls more quickly and with fewer misjudgments. However, ambiguous labeling in situations of vocal ambiguity might have selective advantages. By calling to and clasping an unreceptive female, a male may, over the course of hours, induce her to become sexually receptive. The scarcity of ovipositing females on any given night during the breeding season could heighten the selective pressure on males to locate a female being clasped by another male, even when her calls are ambiguous. Perhaps accuracy could be sacrificed because refraining from advertising to a receptive female has more detrimental consequences than hazarding to pursue an unreceptive female. The energetic cost of mistakenly calling to a ticking female is presumably low, because males call in isolation for hours. Calling has the additional benefits of suppressing other males and possibly attracting another female.
Categorical perception has been described in the perception of
communication and predator signals in a variety of species including crickets,
birds and humans (Liberman et al.,
1957; Liberman et al.,
1961; Nelson and Marler,
1989; Wyttenbach et al.,
1996). It is a perceptual strategy suited to situations in which
identification of signals is more important than fine parameter
discrimination, such as the Polynesian field cricket's identification of
conspecific mating calls and predator bat echolocation calls, which differ
along a spectral continuum (Wyttenbach et
al., 1996). Continuous perception, however, has been described in
the discrimination of intermediate synthetic calls by female green tree frogs:
a two-choice task revealed that auditory resolution is finer than necessary to
distinguish between the principal modulation durations in discrete
vocalizations (Gerhardt, 1978).
Our results suggest that recognition of female calls by male X.
laevis combines elements of both strategies: no discrimination within the
ticking category and continuous discrimination across the overlapping range of
rapping and ticking ICIs.
The ability of the male to discriminate different ICIs (and to categorize
different intervals as belonging to the same class) must rely on the
processing of temporal information within the central nervous system. Our
electrophysiological recordings from auditory nerve fibers and cells in the
first auditory nucleus, the dorsal medullary nucleus, reveal that all click
rates within the vocal range are represented phasically, using a temporal code
of synchronization to the envelope
(Elliott, 2007). However, some
cells in the auditory midbrain (the torus semicircularis) select for click
rate, responding only to certain click rates, using an average spike rate code
(Elliott, 2007). How tuning in
the midbrain contributes to the differential behavioral responses of the male
to female calls is a neuroethological question whose exploration can now be
related to a more complete understanding of the male's perception of
behaviorally relevant temporal cues.
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Acknowledgments |
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References |
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