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The medial olivocochlear (MOC) pathway modulates basilar membrane motion and auditory nerve activity on both a fast (10–100 ms) and a slow (10–100 s) time scale in guinea pigs. The slow MOC modulation of cochlear activity is postulated to aide in protection against acoustic trauma. However in humans, the existence and functional roles of slow MOC effects remain unexplored.
Methodology/Principal Findings
By employing contralateral noise at moderate to high levels (68 and 83 dB SPL) as an MOC reflex elicitor, and spontaneous otoacoustic emissions (SOAEs) as a non-invasive probe of the cochlea, we demonstrated MOC modulation of human cochlear output both on a fast and a slow time scale, analogous to the fast and slow MOC efferent effects observed on basilar membrane vibration and auditory nerve activity in guinea pigs. The magnitude of slow effects was minimal compared with that of fast effects. Consistent with basilar membrane and auditory nerve activity data, SOAE level was reduced by both fast and slow MOC effects, whereas SOAE frequency was elevated by fast and reduced by slow MOC effects. The magnitudes of fast and slow effects on SOAE level were positively correlated.
Citation: Zhao W, Dhar S (2011) Fast and Slow Effects of Medial Olivocochlear Efferent Activity in Humans. PLoS ONE 6(4): e18725. doi:10.1371/journal.pone.0018725
Editor: Izumi Sugihara, Tokyo Medical and Dental University, Japan
Received: November 24, 2010; Accepted: March 9, 2011; Published: April 8, 2011
Copyright: © 2011 Zhao, Dhar. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: This research was funded by grant number DC008420 from the National Institute on Deafness and Other Communication Disorders, and by Northwestern University. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist.
The medial olivocochlear (MOC) pathway, a part of the auditory efferent system, has gained increasing attention in recent years. Its postulated functional roles include protection against acoustic trauma and facilitation of transient-sound perception in a noisy background [1], [2]. Projecting from the medial region of the superior olivary complex, the MOC fibers innervate cochlear outer hair cells (OHCs) via cholinergic synapses. The MOC modulation of the cochlea operates on two time scales, three orders of magnitude apart. Activation of the MOC fibers opens the postsynaptic acetylcholine-gated α9/α10 channels [3], [4], leading to a calcium influx into OHCs [5], [6], [7]. Subsequently, a potassium outflow through calcium-activated potassium channels hyperpolarizes OHCs and decreases the gain of the cochlear amplifier. Hence on a fast time scale (10–100 ms), basilar membrane motion is inhibited and auditory nerve activity reduced. The MOC activity on a slow time scale (10–100 s), although not extensively studied, has been linked to the slow calcium release from intracellular stores and the decrease in OHC's axial stiffness [8], [9], [10]. Both fast and slow MOC effects reduce auditory nerve activity and basilar membrane vibration amplitude in guinea pigs [8], [11], [12]. The fast and slow MOC effects on the phase of basilar membrane displacement, however, are in opposite directions: fast effects causing phase leads and slow effects, phase lags [8]. Furthermore, slow effects peak at a higher frequency than fast MOC effects [12].
MOC effects on the auditory periphery can also be observed using a non-invasive, albeit indirect, method via otoacoustic emissions (OAEs). Spontaneous OAEs (SOAEs) are sounds generated in the cochlea without external stimulation [13]. Their generation is modeled either by standing-wave resonance in the cochlea [13], or by active autonomous oscillation of the stereocilia [18]. Elicited acoustically, the MOC reflex reduces SOAE level and increases its frequency [19], [20], [21], [22], [23] on a time scale consistent with the fast MOC effects observed on basilar membrane mechanics. However, studies in humans have yet to distinguish slow from fast MOC effects.
The present study demonstrates the presence of slow MOC effects on SOAEs in humans. The magnitude of slow MOC effects was miniscule in comparison to that of fast effects.
Thirteen human subjects (ten female and three male), between the ages of 20 and 30 years with normal hearing sensitivity (20 dB HL or better at octave frequencies between 250–8000 Hz, measured with an Interacoustics Audio Traveller AA220) in both ears were recruited for this study. Experiments were conducted on one ear per subject (ten right and three left ears). All subjects selected for the experiments had at least one SOAE 10 dB above the noise floor. All procedures were approved by the Northwestern University Institutional Review Board (Northwestern IRB #4X - Panel E, registration number IRB00000736). Written, informed consent was obtained from each subject. Measurements were conducted in a sound-treated audiological test booth.
Signal generation and recording
Digital stimuli (sampling rate 44100 Hz, 24 bit) were generated by a Macintosh computer and converted to analog signals by a MOTU 828 MKII input/output device. Stimuli were presented to subjects via MB Quart 13.01 HX transducers coupled to the ear canal with an Etymotic Research ER10B+ probe. OAE signals were acquired in subjects' ear canals using the ER10B+ microphone, amplified by a pre-amplifier (+20 dB), digitized by the MOTU and stored on disk for processing offline.
Fast and slow MOC effects on SOAEs
MOC efferents were activated by a contralateral broadband noise (100–10000 Hz, 68 dB SPL, 5 ms rise/fall time). A higher-level (83 dB SPL) contralateral noise was applied to three subjects in a subset of the experiments. Each run consisted of a pre-stimulation window (50 s), a stimulation window (102 s) and a post-stimulation window (150 s) (Figure 1). As an acoustical approximation of the stimulation paradigm employed by Sridhar et al. (1995) and Cooper and Guinan (2003) [8], [12], 3-s noise bursts were presented in the contralateral ear interleaved with 3-s silent intervals within the stimulation window. A total of six runs were recorded and a zero-padded fast Fourier transform (FFT) was performed for each 1-s window to generate SOAE spectra with a frequency resolution of 0.5 Hz. The noise floor of each 1-s window was taken as the median spectral level in a 100-Hz range surrounding the target SOAE. SOAE data were rejected off-line if SOAE level in the local time window was less than two standard deviations above the average noise floor for the entire recording.
Each SOAE was monitored for 302 s. Baseline SOAE level and frequency were established in the pre-stimulation window. Contralateral broadband noise pulses (3-s long, 68 dB SPL) were presented in the 102-s stimulation window with 3-s inter-pulse intervals. Blue arrows represent SOAE estimates in the pre- and post-stimulation windows, red arrows during noise pulses, and green arrows during inter-pulse intervals. This color convention is consistent throughout the paper. Each arrow represents an estimate of SOAE level or frequency averaged over a 1-s window.
Middle-ear reflex test
The middle-ear muscle (MEM) and MOC reflexes share the same afferent pathway and are elicited by external sounds of similar levels. Hence, it is critical to ensure the MEM reflex remains dormant while the MOC reflex is elicited. Two experimental methods, the group delay method [24] and the suppression method [25], were implemented in all subjects to monitor the MEM reflex. Both methods utilize stimulus frequency OAEs (SFOAEs), which are low-level acoustic signals evoked by a single-frequency tone and measured in the ear canal [26]. SFOAEs can be extracted by nonlinear compression, two-tone suppression, or spectral smoothing [27]. Both suppression and compression techniques were used in our MEM contraction test to extract SFOAEs evoked by a 40 dB SPL probe tone.
In the group delay method, the vector difference between the total ear canal pressure at the probe frequency with and without contralateral noise was denoted ΔP. The group delay of ΔP was computed as the negative of its phase slope. Around 1500 Hz, a ΔP group delay near 10 ms indicates the dominance of the MOC reflex over the MEM reflex [24], [28]. The group delay of ΔP was measured using both a sweeping probe tone paradigm and a discrete probe tone paradigm (Figure S1), as described below. In the sweeping probe tone paradigm, a probe was swept from 800 to 2400 Hz in 12 s. The following triplet was repeated eight times: a 40 dB SPL probe in the absence of contralateral noise, followed by a 60 dB SPL probe in the absence of contralateral noise, and finally a 40 dB SPL probe paired with contralateral noise. Averaged data were passed through an adaptive least-squares fit filter to estimate the level and phase of the total ear canal pressure at the probe frequency [29]. Baseline SFOAE was calculated using the presumed nonlinear compressive growth of SFOAEs [26]. ΔP was also calculated as described above. In the discrete probe paradigm, a discrete tone at 40 dB SPL was presented over an 80-Hz range in 20-Hz steps. The probe tone lasted 5 s, during which time a 68 dB SPL contralateral noise was presented from 0.5 to 3 s. A total of eight runs were performed for each probe frequency and averaged data were passed through an adaptive least-squares-fit filter to obtain probe level and phase.
In the suppression method, SFOAEs were evoked by a 40 dB SPL probe tone at three frequencies (602 Hz, ∼1000 Hz, ∼2000 Hz) in each subject. The probe frequencies around 1000 Hz and 2000 Hz were chosen to be at a local peak of the ΔP versus probe frequency function obtained using the sweeping tone paradigm described earlier. Each recording block in this paradigm lasted 12.5 s (Figure S2A) and was divided into five segments. Four such blocks were averaged to obtain usable signal-to-noise ratios. A probe tone (P) was presented during the entire 12.5 s; a 65 dB SPL suppressor tone (S), 0.1 octaves below the probe, was presented between 2.5 and 7.5 s; and a contralateral broadband noise (C) was presented between 5 and 10.5 s. Thus, the resulting five segments were: probe alone (P), probe plus suppressor (P+S), probe plus suppressor and contralateral noise (P+S+C), probe plus contralateral noise (P+C), and probe alone again (P′). The total ear canal pressure at the probe frequency was measured for each of the five segments. The vector difference between P and P+S yields baseline SFOAE , between P+S and P+S+C yields the pressure change due to potential MEM contraction (blue arrow, Figure S2B), between P and P+C yields ΔP (red arrow, Figure S2B), and between P and P′ is a measure of drift in our measurements (green arrow, Figure S2B). The SFOAE is arguably largely suppressed in the P+S window. Therefore, any differences between the complex pressures in the ear canal measured in the P+S and P+S+C segments can be attributed to the MEM reflex [25]. When the magnitude of ΔP is substantially larger than that of, and therefore cannot be explained by either MEM reflex-induced pressure change or probe drift, ΔP is considered to be dominated by the MOC reflex. For two out of three probe frequencies (∼1000 Hz and ∼2000 Hz), the exemplar subject in Figure S2C displayed a ΔP (red symbols) that was substantially larger than both pressure change by MEM reflex (blue symbols) and probe drift (green symbols), indicating the dominance of the MOC reflex.
Using the above two methods, we ensured the absence of MEM contraction under a 68 dB SPL contralateral noise.
Effects of contralateral noise on forty-four SOAEs from thirteen subjects, between 871 and 14864 Hz in frequency, and −7 and 18 dB SPL in level, were examined on a fast time scale (evaluated during 3-s bursts of contralateral noise) and on a slow time scale (measured in a 30-s window after noise stimulation). SOAE levels were reduced on both the fast and the slow time scale. SOAE frequencies were elevated on the fast time scale but reduced on the slow time scale. The magnitude of fast effects was significantly greater than that of slow effects.
Demonstration of MOC-induced changes in SOAEs
An example of MOC-induced changes in SOAE level and frequency is presented in Figure 2 (SOAE at 1702 Hz/16 dB SPL, subject WTPF01). Six consecutive trials of the 302-s noise stimulation paradigm (represented by the black rectangular box in Figure 2A), totaling 1812 s, were recorded from each subject. SOAE level and frequency throughout the entire 1812 s are shown in Figure 2A and 2C, respectively. SOAE level and frequency averaged over six runs are presented in Figure 2B and 2D. Blue symbols depict estimates of SOAE level (Figure 2A, 2B) and frequency (Figure 2C, 2D) in the pre- and post-stimulation windows. Estimates of SOAE level and frequency during and between noise pulses are presented using red and green symbols, respectively.
SOAE level over six consecutive repetitions of the stimulation paradigm (A), and averaged level over six repetitions (C) are shown in the top row (blue: pre- and post-stimulation windows; red: during noise pulses; green: inter-pulse intervals). The black rectangular box in Panel A represent the noise pulses which are shown only for the first experimental run but were repeated six times. Baseline () is defined as the median SOAE level/frequency in the 30-s window before noise onset. Similarly, SOAE levels/frequencies during the first two noise pulses (), during the last two noise pulses (), during the last two inter-pulse intervals () and in the 30-s window after noise offset () were quantified. Differences from the baseline were defined as fast change (-), adaptation change (-), buildup change (-) and slow change (-). Error bars represent one standard deviation.
Baseline values of SOAE level and frequency were established by averaging these values over the 30-s window before noise onset ( in Figure 2B, 2D). Noise pulses suppressed SOAE level (red symbols, Figure 2A, 2B) and elevated SOAE frequency (red symbols, Figure 2C, 2D). Differences between the estimates of SOAE level and frequency during the first two noise pulses (0–3 & 6–9 s of the stimulation window, in Figure 2B, 2D) and the baseline were defined as fast changes (-). Fast changes represent fast MOC effects as the measurement epoch was limited to the first few seconds of the stimulation window, thereby eliminating the influence of slow MOC effects. As evident in Figure 2B and 2D, changes in both SOAE level and frequency gradually adapted over tens of seconds during noise pulses. Differences between SOAE estimates during the last two noise pulses (90–93 & 96–99 s of the stimulation window, in Figure 2B, 2D) and the baseline were defined as adaptation changes (-). This metric reflects the mixed effects of adapted fast and slow MOC effects. Residual SOAE changes in the inter-pulse intervals and after noise offset were much smaller than changes during noise pulses. Differences between estimates of SOAE level and frequency during the last two inter-pulse intervals (93–96 & 99–102 s of the stimulation window, in Figure 2B, 2D) and the baseline were defined as buildup changes (-). Buildup changes are free of fast MOC effects, but manifest a mixture of slow MOC effects and a post-noise overshoot [23]. Differences between averaged SOAE estimates in the first 30 s of the post-stimulation window ( in Figure 2B, 2D) and the baseline were defined as slow changes (-). Slow changes are a pure representation of slow MOC effects, as both fast MOC effects and the post-noise overshoot dissipate in the first few seconds of the post-stimulation window. Both buildup and slow changes were smaller in magnitude than fast and adaptation changes. Group data are presented below.
Group Results
Fast, adaptation, buildup and slow changes were quantified for individual SOAEs (N = 44) and are presented in Figure 3. SOAE level changes in dB (left column, Figure 3), SOAE frequency changes in Hz (middle column, Figure 3) and SOAE frequency change as percentage of baseline frequency (right column, Figure 3) are plotted as functions of baseline SOAE level (upper row, Figure 3) and frequency (lower row, Figure 3). Filled and open symbols represent statistically significant (t-test, p<0.01) and non-significant (t-test, p≥0.01) changes from baseline, respectively. This convention to distinguish significant from non-significant changes is preserved throughout the paper. The grand averages for fast change (N = 32), adaptation change (N = 33), buildup change (N = 44) and slow change (N = 44) are presented in Figure 4. The analysis exclusion criterion (excluding SOAEs less than two standard deviation above the local noise floor) accounts for the discrepancy between the overall sample size (N = 44) and the sample sizes of fast change (N = 32) and adaptation change (N = 33).
Filled and open symbols represent statistically significant (t-test, p<0.01) and non-significant (t-test, p≥0.01) changes from baseline, respectively. Fast and adaptation changes are larger in magnitude than buildup and slow changes.
The results of a paired Wilcoxon signed-rank test demonstrated that slow and buildup changes were significantly smaller than fast and adaptation changes (*: p<1e-6). The central line on each box is the median value. The top and bottom edge lines represent 25th and 75th percentiles, respectively. Whiskers cover all data points within 1.5 interquartile range from the top and bottom edge lines. Red crosses mark outliers beyond the whiskers.
Fast and adaptation changes manifested as decrease in SOAE level and elevation in SOAE frequency (Wilcoxon signed-rank test, p<1e-6) (Figures 3 and 4). The median fast changes were −5.3 dB in level and 10.7 Hz in frequency (0.32% of baseline frequency). The median adaptation changes were −3.6 dB in level and 9.3 Hz in frequency (0.28% of baseline frequency). Note that we implemented an analysis exclusion criterion, which rejects SOAEs less than two standard deviations above the local noise floor. Since low-level SOAEs are more likely to be suppressed into or near the noise floor and thus be excluded from analyses, fast and adaptation changes as functions of baseline SOAE level (upper row, Figure 3) could have been tainted by this exclusion criterion, which probably accounts for the trend that SOAEs below 5 dB SPL had less fast and adaptation changes in level than larger SOAEs (Figure 3A). No prominent feature was observed in the fast and adaptation changes versus baseline SOAE level functions. Fast and adaptation changes as functions of baseline SOAE frequency (lower row, Figure 3) however demonstrated clear patterns. Fast and adaptation changes in both level (in dB) and frequency (as percentage of baseline frequency) were greater for low- to mid-frequency SOAEs than for high-frequency SOAEs (Figure 3D and 3F).
Buildup and slow changes manifested as reductions in both SOAE level and frequency (Wilcoxon signed-rank test, p<0.01) (Figures 3 and 4). The median buildup changes were −0.2 dB in level and −0.6 Hz in frequency (−0.02% of baseline frequency). The median slow changes were −0.2 dB in level and −0.2 Hz in frequency (−0.01% of baseline frequency). No dependence of buildup or slow changes on either baseline level or frequency could be easily identified.
The paired Wilcoxon signed-rank test was performed to compare the four types of changes. Different ears emitted different numbers of SOAEs at diverse frequencies and levels, and each SOAE was treated as an independent observation. Fast and adaptation changes were significantly larger in magnitude than buildup and slow changes (Paired Wilcoxon signed-rank test, p<1e-6) (Figure 4).
To demonstrate the relationship between fast and slow changes, fast changes are plotted against slow changes (N = 32) (Figure 5). Filled symbols indicate that both fast and slow changes were statistically significant (t-test, p<0.01), whereas open symbols indicate otherwise. A weak positive correlation (r = 0.46, p<0.01) was observed between fast and slow changes in SOAE level (both filled and open symbols included), consistent with a sequential relationship in a series of physiological events. Correlation between fast and slow changes in SOAE frequency was not statistically significant (p>0.05).
Filled symbols indicate that both fast and slow changes were statistically significant (t-test, p<0.01), whereas open symbols indicate otherwise. A positive correlation for SOAE level was observed. Dashed lines represent best linear fits to all of the data points.
Effect of contralateral noise level on slow changes
The contralateral noise level of 68 dB SPL was selected for our experiments in order to avoid elicitation of the MEM reflex. This mild noise level may have led to insufficient activation of the MOC bundle, resulting in the relatively small slow changes in SOAEs (Figures 3 and 4). To study the influence of contralateral noise level on the magnitude of slow changes, we repeated the experiments in three subjects on a different day with a higher noise level of 83 dB SPL. The slow changes were evaluated over a 30-s post-stimulation window while the potential MEM reflex has a time constant in the range of hundreds of milliseconds [30]. Given these differences in the time courses between slow MOC effects and the MEM reflex, MEM contraction, if elicited by the 83 dB SPL noise, arguably had little influence on the slow changes.
SOAEs that were present on both days were examined in detail (N = 11). Increasing noise level from 68 to 83 dB SPL did not substantially enhance the magnitude of slow changes (Figure 6). Filled symbols indicate slow changes elicited by both noise levels were statistically significant (t-test, p<0.01), whereas open symbols indicate otherwise. Data points cluster along the diagonal, suggesting no increase in the magnitude of slow effects as the noise level increased from 68 to 83 dB SPL. The reader is reminded that the changes should be evaluated in their absolute values (i.e., a value of −2 dB is a greater change than a value of −1 dB).
Filled symbols indicate slow changes elicited by both noise levels were statistically significant (t-test, p<0.01), whereas open symbols indicate otherwise. No prominent elicitor-level effect on slow changes was observed.
The discovery of MOC modulation of cochlear output on a slow time scale (10–100 s) in guinea pigs prompted speculations on its functional roles in protection against acoustic trauma [10], [12], [31]. Here we demonstrate that contralateral noise (68 and 83 dB SPL) elicits miniscule yet significant slow MOC effects on SOAEs in human subjects, consistent with a high threshold or small magnitude of slow MOC effects.
Interference from MEM contraction
Since the MOC reflex and the MEM reflex share similar temporal characteristics, and can both be elicited by contralateral broadband noise, it is difficult to parse the modulation of OAEs by these two reflexes. In this work, absence of MEM contraction when applying a 68 dB SPL contralateral noise was ensured using two experimental methods, the group delay method and the suppression method. In the group delay method, the group delay of ΔP induced by contralateral noise was around 10 ms in all subjects (Figure S1). In the suppression method, three probe frequencies were applied (602 Hz, ∼1000 Hz and ∼2000 Hz), and ΔP was substantially larger than the pressure change caused by MEM reflex or probe drift for at least two of the three probe frequencies (Figure S2). Nonetheless, substantial probe drift or the minimal magnitude of ΔP prevented unambiguous exclusion of the MEM reflex at some probe frequencies. Overall, these controls made us reasonably confident that our findings were not strongly influenced by MEM contraction. Thus, the slow modulation of SOAEs can be attributed to slow effects of the MOC efferents.
Fast and slow MOC effects on human SOAEs
Both fast and slow MOC effects led to a reduction of SOAE level in humans (Figure 4A). Consistent with human data, both fast and slow MOC effects in guinea pigs reduce basilar membrane motion and auditory nerve activity [8], [11], [12]. The reduction in all these cochlear measures stems from the MOC-induced attenuation of the gain of the cochlear amplifier.
Fast MOC effects led to an elevation in SOAE frequency whereas slow MOC effects cause a reduction (although small) in SOAE frequency (Figure 4B and 4C). In guinea pigs, fast and slow MOC effects have been shown to produce phase changes of basilar membrane motion in opposite directions, fast effects producing phase leads and slow effects phase lags [8], [32]. Congruity between OAE and basilar membrane data can be achieved under the framework of the global standing wave model of SOAEs [15], where phase leads in basilar membrane vibration predict elevation in SOAE frequency, and phase lags predict decrease in SOAE frequency [23].
The magnitude of fast MOC effects on both SOAE level and frequency (as percentage of baseline frequency) appears to diminish gradually above 6 kHz with increasing frequency (Figure 3D and 3F). This is in agreement with the frequency dependence of fast MOC effects in guinea pigs, which drops off above 10 kHz [12].
Comparing slow MOC effects between humans and guinea pigs
In guinea pigs, the magnitudes of slow and fast MOC effects are not drastically different. The magnitude of the slow MOC suppression of auditory nerve activity peaks at a higher frequency (∼16 kHz) compared to that of fast MOC suppression (6–10 kHz), and the peak magnitude of slow effects is three to four times smaller than that of fast effects [11], [12]. Fast and slow MOC effects on basilar membrane vibration are also comparable in size [8].
In contrast to data obtained from guinea pigs, slow MOC effects on human SOAEs are strikingly small compared to fast MOC effects (Figure 4). The median slow effects were −0.2 dB (level change) and −0.2 Hz (frequency change), whereas the median fast effects were −5.3 dB and 10.7 Hz.
Although we highlight the miniscule magnitude of slow MOC effects in our data, we did not fully account for their frequency dependence. While slow effects peak at high frequencies (∼16 kHz) in guinea pigs [12], the propensity of measurable SOAEs to cluster in the low frequency range (1–4 kHz) in humans limited our scope. However, it bears mention that slow changes in SOAE level or frequency were minimal for SOAEs above ∼6 kHz (Figure 3B, 3D). The lack of pronounced slow effects across all frequencies evaluated in humans may be due to differences in species and in experimental design (e.g., stimulus, stimulation paradigm, measures, etc.).
Another critical difference between our experiments and those in guinea pigs is the intensity of MOC efferent stimulation. The shock rate of 150/s used in the continuous paradigm of Sridhar et al. (1995) is roughly equivalent to acoustic stimulation over 90 dB SPL [12]. Hence, our 68 dB SPL contralateral noise is much weaker an elicitor of efferent activity than electrical shocks delivered to guinea pigs. To partially address this difference in elicitor intensity, we examined three subjects with a higher-level contralateral noise at 83 dB SPL but observed no substantial increase in the magnitude of slow effects (Figure 6).
Arguably, the slow effects in this study are qualitatively similar to those in guinea pigs. With a stimulation paradigm that was an acoustical approximation of that employed by Sridhar et al. (1995) and Cooper and Guinan (2003), we observed a positive correlation between fast and slow changes in SOAE level (Figure 5A), consistent with the positive correlation in guinea pigs between fast and slow effects as stimulus level increases [12]. In contrast, Larsen and Liberman (2009) applied lengthened continuous noise stimulation (5-min long) in guinea pigs and reported a negative correlation between 'onset' suppression and 'buildup' suppression of compound action potential [33]. In mice, electrically shocking the olivocochlear efferents leads to a post-shock enhancement of sound-evoked compound action potential and distortion product OAEs over tens or even hundreds of seconds [34]. This novel enhancement, which is independent of α9 cholinergic receptors, probably does not share common underlying mechanisms as the suppressive slow MOC effects observed in guinea pigs and humans, which reduce the amplitude of cochlear output.
The auditory efferents are thought to play a protective role against acoustic trauma. This protective role has been associated with slow MOC effects elicited by electrical shocks in guinea pigs [31]. In these experiments, MOC efferents were stimulated by the electrical equivalent of noise over 90 dB SPL. We have shown that contralateral noise up to 83 dB SPL elicited miniscule slow MOC effects in humans. However, our results did not rule out a higher threshold for larger slow effects or rapid growth in the magnitude of slow effects with increasing noise level. In three human subjects, the magnitude of slow effects did not grow dramatically as the level of contralateral noise increased from 68 to 83 dB SPL. It is yet to be seen whether noise over 90 dB SPL produces prominent slow effects. The functional relevance of slow MOC effects remains elusive.
Supporting Information
Figure S1.
Example of the group delay method for detecting middle-ear muscle (MEM) contraction. A 40 dB SPL probe tone was either swept across, or presented at discrete frequencies in 20-Hz steps over an 80-Hz range near 1000 Hz. The total ear canal pressure was measured with and without a 68 dB SPL contralateral noise. The magnitude (A) and phase (B) of the vector difference in the total ear canal pressure at the probe frequency between the two conditions, denoted ΔP, was plotted as a function of probe frequency. A group delay of ΔP around 10 ms indicates the dominance of the MOC reflex over the MEM reflex.
Figure S2.
The suppression method for detecting MEM contraction. (A) Illustration of the experimental paradigm. A 40 dB SPL probe tone, a 65 dB SPL suppressor tone 0.1 octave below the probe tone, and a 68 dB SPL contralateral noise were presented for different durations in a 12.5-s window, segmenting it into five conditions: probe alone (P), probe plus suppressor (P+S), probe plus suppressor and contralateral noise (P+S+C), probe plus contralateral noise (P+C), and finally probe alone again (P′). Computing the vector difference between these conditions yields baseline SFOAE, pressure change induced by the middle-ear muscle reflex (MEMR) (blue arrow), contralateral noise-induced shift ΔP (red arrow) and probe drift (green arrow) (B). Exemplar results from subject WTPF42 are displayed (C). For two SFOAE probe frequencies (∼1000 and ∼2000 Hz), the magnitude of ΔP (red symbols) was substantially larger than that of, and therefore could not be explained by either MEMR-induced pressure change (blue symbols) or probe drift (green symbols). Hence ΔP was considered to be dominated by the MOC reflex. Error bars are one standard error.
The authors would like to thank Jonathan Siegel, Christopher Shera, Claus-Peter Richter, Steven Zecker and Jungwha Lee for stimulating discussions, and two anonymous reviewers for constructive comments. We are also grateful to Rachael Baiduc, James Dewey, Kate Dunckley, Jungmee Lee and Gayla Poling for comments on the manuscript, and Dashiell Oatman-Stanford for his involvement in pilot data collection. Parts of this work were presented at the 2010 annual convention of the Association for Research in Otolaryngology.
Author Contributions
Conceived and designed the experiments: WZ SD. Performed the experiments: WZ. Analyzed the data: WZ SD. Wrote the paper: WZ SD.
- 1. Guinan JJ Jr (2006) Olivocochlear efferents: anatomy, physiology, function, and the measurement of efferent effects in humans. Ear Hear 27: 589–607.
- 2.
Robles L, Delano PH (2008) Efferent System. In: Dallos P, Oertel D, editors. The Senses: A Comprehensive Reference. 1 ed. Academic Press.
- 3. Elgoyhen AB, Johnson DS, Boulter J, Vetter DE, Heinemann S (1994) Alpha 9: an acetylcholine receptor with novel pharmacological properties expressed in rat cochlear hair cells. Cell 79: 705–715.
- 4. Elgoyhen AB, Vetter DE, Katz E, Rothlin CV, Heinemann SF, et al. (2001) alpha10: a determinant of nicotinic cholinergic receptor function in mammalian vestibular and cochlear mechanosensory hair cells. Proc Natl Acad Sci USA 98: 3501–3506.
- 5. Fuchs PA, Murrow BW (1992) Cholinergic inhibition of short (outer) hair cells of the chick's cochlea. J Neurosci 12: 800–809.
- 6. Fuchs PA, Murrow BW (1992) A novel cholinergic receptor mediates inhibition of chick cochlear hair cells. Proc Biol Sci 248: 35–40.
- 7. Housley GD, Ashmore JF (1991) Direct measurement of the action of acetylcholine on isolated outer hair cells of the guinea pig cochlea. Proc Biol Sci 244: 161–167.
- 8. Cooper NP, Guinan JJ Jr (2003) Separate mechanical processes underlie fast and slow effects of medial olivocochlear efferent activity. J Physiol 548: 307–312.
- 9. Dallos P, He DZ, Lin X, Sziklai I, Mehta S, et al. (1997) Acetylcholine, outer hair cell electromotility, and the cochlear amplifier. J Neurosci 17: 2212–2226.
- 10. Sridhar TS, Brown MC, Sewell WF (1997) Unique postsynaptic signaling at the hair cell efferent synapse permits calcium to evoke changes on two time scales. J Neurosci 17: 428–437.
- 11. da Costa DL, Chibois A, Erre JP, Blanchet C, de Sauvage RC, et al. (1997) Fast, slow, and steady-state effects of contralateral acoustic activation of the medial olivocochlear efferent system in awake guinea pigs: action of gentamicin. J Neurophysiol 78: 1826–1836.
- 12. Sridhar TS, Liberman MC, Brown MC, Sewell WF (1995) A novel cholinergic "slow effect" of efferent stimulation on cochlear potentials in the guinea pig. J Neurosci 15: 3667–3678.
- 13. Kemp DT (1979) Evidence of mechanical nonlinearity and frequency selective wave amplification in the cochlea. Arch Otorhinolaryngol 224: 37–45.
- 14. Kemp DT (1979) The evoked cochlear mechanical response and the auditory microstructure - evidence for a new element in cochlear mechanics. Scand Audiol Suppl: 35–47.
- 15. Shera CA (2003) Mammalian spontaneous otoacoustic emissions are amplitude-stabilized cochlear standing waves. J Acoust Soc Am 114: 244–262.
- 16. Talmadge CL, Tubis A, Long GR, Piskorski P (1998) Modeling otoacoustic emission and hearing threshold fine structures. J Acoust Soc Am 104: 1517–1543.
- 17. Zweig G, Shera CA (1995) The origin of periodicity in the spectrum of evoked otoacoustic emissions. J Acoust Soc Am 98: 2018–2047.
- 18. Martin P, Hudspeth AJ (1999) Active hair-bundle movements can amplify a hair cell's response to oscillatory mechanical stimuli. Proc Natl Acad Sci U S A 96: 14306–14311.
- 19. Harrison WA, Burns EM (1993) Effects of contralateral acoustic stimulation on spontaneous otoacoustic emissions. J Acoust Soc Am 94: 2649–2658.
- 20. Mott JB, Norton SJ, Neely ST, Warr WB (1989) Changes in spontaneous otoacoustic emissions produced by acoustic stimulation of the contralateral ear. Hear Res 38: 229–242.
- 21. Rabinowitz WM, Widin GP (1984) Interaction of spontaneous oto-acoustic emissions and external sounds. J Acoust Soc Am 76: 1713–1720.
- 22. Schloth E, Zwicker E (1983) Mechanical and acoustical influences on spontaneous oto-acoustic emissions. Hear Res 11: 285–293.
- 23. Zhao W, Dhar S (2010) The effect of contralateral acoustic stimulation on spontaneous otoacoustic emissions. J Assoc Res Otolaryngol 11: 53–67.
- 24. Guinan JJ Jr, Backus BC, Lilaonitkul W, Aharonson V (2003) Medial olivocochlear efferent reflex in humans: otoacoustic emission (OAE) measurement issues and the advantages of stimulus frequency OAEs. J Assoc Res Otolaryngol 4: 521–540.
- 25. Lilaonitkul W, Guinan JJ Jr (2009) Reflex control of the human inner ear: a half-octave offset in medial efferent feedback that is consistent with an efferent role in the control of masking. J Neurophysiol 101: 1394–1406.
- 26.
Kemp DT, Chum RA (1980) Observations of the generator mechanism of stimulus frequency acoustic emissons. In: van den Brink G, Bilsen FA, editors. Psychophysical, Physiological and Behavioural Studies in Hearing. Delft, The Netherlands: Delft University Press. pp. 34–42.
- 27. Kalluri R, Shera CA (2007) Comparing stimulus-frequency otoacoustic emissions measured by compression, suppression, and spectral smoothing. J Acoust Soc Am 122: 3562–3575.
- 28. Shera CA, Guinan JJ Jr (1999) Evoked otoacoustic emissions arise by two fundamentally different mechanisms: a taxonomy for mammalian OAEs. J Acoust Soc Am 105: 782–798.
- 29. Long GR, Talmadge CL (1997) Spontaneous otoacoustic emission frequency is modulated by heartbeat. J Acoust Soc Am 102: 2831–2848.
- 30. Goodman SS, Keefe DH (2006) Simultaneous measurement of noise-activated middle-ear muscle reflex and stimulus frequency otoacoustic emissions. J Assoc Res Otolaryngol 7: 125–139.
- 31. Reiter ER, Liberman MC (1995) Efferent-mediated protection from acoustic overexposure: relation to slow effects of olivocochlear stimulation. J Neurophysiol 73: 506–514.
- 32. Murugasu E, Russell IJ (1996) The effect of efferent stimulation on basilar membrane displacement in the basal turn of the guinea pig cochlea. J Neurosci 16: 325–332.
- 33. Larsen E, Liberman MC (2009) Slow build-up of cochlear suppression during sustained contralateral noise: central modulation of olivocochlear efferents? Hear Res 256: 1–10.
- 34. Maison SF, Vetter DE, Liberman MC (2007) A novel effect of cochlear efferents: in vivo response enhancement does not require alpha9 cholinergic receptors. J Neurophysiol 97: 3269–3278.


text:
The first Wagamama opened close to central London's Tottenham Court Road in April, 1992; the second, in nearby Lexington Street, started just over a year ago. But the restaurants have been so effective at capturing the public imagination that the operation already has a turnover of about pounds 5.5m.
Foreign & Colonial Ventures last year took a 33 per cent stake in the business, and 34-year-old Mr Yau is confident that he can open a third location this year. He has plans to open four more sites in London over the next three years and branch out by setting up in Manchester and other regional centres. Once there are six to eight units, he thinks the company "should be geared up for some sort of flotation".
And after that? "I think our aim is really to take the concept to the United States. The concept is made for the US market. Potentially, it could be biggest there," he says.
That concept is a niche between fast food and casual dining. The idea is that customers go to a Wagamama on the way to somewhere else, such as the cinema or a tourist attraction. Moreover, the cooking - a Japanese cuisine known as Chinese-style noodles - is easy to master for non-Orientals and caters more easily to these health-conscious times than burgers and other fast foods.
Though customers share long tables and sit on simple benches, the food is served at the table rather than collected. And, with his keen eye for detail, Mr Yau explains that the typical customer is aged between 18 and 35. During the day, he or she is likely to be a tourist or office worker, but less easily definable in the evening. The average spend is pounds 9 and the average stay is half an hour during the day and an hour in the evening.
Although Mr Yau's father was sceptical about something as untried as noodles for the mass market, he was one of the original investors. He made the right decision: the business was serving 1,000 customers a day in the first year, and he made a good return when Foreign & Colonial bought out him and the other shareholders.
In fact, the success was so great that the company soon had problems obtaining enough noodles to meet demand. Having started with a local supplier, it had to resort to importing the basis of its menu direct from Japan - which created its own problems when the pound started to fluctuate against the yen.
Though Mr Yau says that dealing with this was harder than running the business, he has so far resisted bringing in much in the way of outside management. He is planning to recruit a non-executive chairman to raise the company's profile in the run-up to flotation, but has largely relied on the advice of Rees Pollock, a small accountancy firm that specialises in helping growing businesses.
His confidence that the idea has international appeal comes from a combination of the interest in healthier eating, comments from his customers and the fact that the majority of the investment enquiries he receives every week are from the US.
"I believe that noodles are going to be the next pizza," he says. "When we first opened there was just us plus one other. Now, there's more than 10 in London."
- More about:
- Mcdonald's
- Noodles
- Restaurants
- Tourism


text:
A Poor Excuse for a Dragon
Random House Children's Books, Aug 23, 2011 - Juvenile Fiction - 48 pages
"Punchy dialogue and compact sentences should make this a read-aloud delight."—Publishers Weekly
"Part-slapstick, part-fairy tale, the gently humorous plot has enough twists and turns to keep newly independent readers engaged."—School Library Journal
Fred the dragon has a list of tasks he must complete in order to be a successful dragon—none of which comes naturally. But he's determined to make #5—eat people—work. Before you can say "pass the salt" he's gobbled up three people even though he doesn't have the stomach for it. Luckily a local shepherd, with the help of a giant and a witch, knows how to cure what ails him and get those pesky people out of his belly. It's happily-ever-after for everyone in ways you'd never expect.
From the Trade Paperback edition.


text:
|WTF Priorities?
|attached to China Rejects US Piracy Claims As "Groundless"
|Re:some fine police officers who don't deserve
|attached to Writer Peter Watts Sentenced; No Jail Time
|Re:I caught several cheaters
|attached to How Easy Is It To Cheat In CS?
|Re:A comment from Tynt
|Re:A comment from Tynt
|Re:A comment from Tynt
|Re:A comment from Tynt
|attached to Tynt Insight Is Watching You Cut and Paste
|Re:Everyone has their price?
|attached to Mark Cuban's Plan To Kill Google
|Re:they've been copying Mac all along...
|attached to Microsoft Responds To "Like OS X" Comment
|Re:Don't make the pair programming compulsory
|attached to Collaborative Software For Pair Programming?
|attached to Cure For Radiation Sickness Found?
|Re:I know why.
|Re:I know why.
|attached to Bill Gates Puts Classic Feynman Lectures Online
|Re:Mutually Assured Destruction? I think not...
|attached to Microsoft vs. Google — Mutually Assured Destruction
|Re:No! Don't tell us!
|attached to Military's Satellite Meteor Data Sharing May Soon Resume
|Re:The law is on London's side
|attached to UK's National Portrait Gallery Threatens To Sue Wikipedia User
|Re:CORY DOCTOROW IS NOT A "PROF."
|Re:The Professor is an Idiot
|attached to Student Who Released Code From Assignments Accused of Cheating
|Imagine that
|attached to Sony CEO Proposes "Guardrails For the Internet"
|Re:General sections are free, Financial aren't
|Re:Best of luck with that.
|attached to News Corp Will Charge For Newspaper Websites
|Re:Reciprocal regulations
|attached to China Considering Cuts In Rare-Earth Metal Exports
Heuristics are bug ridden by definition. If they didn't have bugs, then they'd be algorithms.


text: Lego's Four-Foot Super Star Destroyer Executor Will Be Longest Official Set Ever SoldJack Loftus6/19/11 7:00pmFiled to: LegoLego Super Star DestroyerDarth VaderStarwarsModels bricks record104EditPromoteShare to KinjaToggle Conversation toolsGo to permalink On September 1, Lego will begin selling Darth Vader's Super Star Destroyer, the Executor. At 50 inches and eight pounds, it will be the longest set Lego has ever sold. Advertisement With the set you'll get Darth Vader and some of those memorable bounty hunters from the original trilogy."Just" $400 and this mega-model is yours on September 1. [CNET]Reply104 repliesLeave a reply


text:
Recent Tenants Harbor Reviews
wonderful experience
Lobster Harbor View was a perfect house for our family. It was clean and airy with plenty of space for all. The kitchen was great and we really used it! We attended a wedding at the East Wind Inn in Tenants Harbor which was within walking distance. The walk was beautiful overlooking the harbor and a nearby General Store very convenient. The owners were very accommodating and easy to work with. Great place, great time! C. Delaney


text:
In a message dated 7/7/01 7:35:49 PM Eastern Daylight Time,
firstname.lastname@example.com writes:
<< Which leads me to: do hosta roots need to go deep? My thinking is
10 inches is all they need. >>
My two year old Blue Angel would need a tree spade to move it. It is
competing with a much older S&S for size.
I think that I will go with the three gallon size and will have to have my
husband figure out how big that would be. Maybe a little more than ten
inches for depth, my hostas in gallon pots seem to fill up the bottom of the
pot if given time.
Thanks for helping me to sort this one out in my head. N.
To sign-off this list, send email to firstname.lastname@example.com with the
message text UNSUBSCRIBE HOSTA-OPEN


text:
Well how about an F-15C?
I'll second that. The lady that owned the place was a class act.
No problem. Keep in in mind though - if nothing else, it makes a great tool to use at home instead of paying through the nose for D3 or Reality when you're just doing local dev.
Why not use ScarletDME? It's a fork of the open source OpenQM project and would handle your needs quite well. There's a couple of ways to access it via PHP as well. If you know Pick no reason not to stay with it.
...is doing the same kind of scammy crap to the FlightGear project. They've taken a very old build and are selling it for an absurd amount of money. Some people have been taken for over $100 with the scam. Clickbank seems to help them out too.
See http://www.flightgear.org/flightprosim.html for more info. If you spot these jerks on Facebook, please report their page as a scam.
Recursion is the root of computation since it trades description for time.


text:
PlayStation Help
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yeah this is where I'm so screwed.... I only have one rifle shot left and about 30 handgun bullets... I can't seem to kill the monster before I run out. Is there a cheat code? because I'll be damned if I'm starting this game over again!
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Climb UP
Oklahoma City, OK 73129
Climb UP, which originated in Norman, took over the old Rocktown Climbing Gym silos in Bricktown, keeping this...Learn More
Climb UP, which originated in Norman, took over the old Rocktown Climbing Gym silos in Bricktown, keeping this...Learn More
Catch some serious air at the outdoor Ardmore Sk8 Park inside the Ardmore Regional Park. Bring your boards...Learn More
Bordered by Fort Sill on its south shores, the Wichita Mountains Wildlife Refuge on its north shores and sitting in the...Learn More
Gene's Dive Shop has been servicing the Lake Tenkiller Area since 1971. Located near the Tenkiller Dive Park,...Learn More
Nautical Adventures Scuba is Lake Tenkiller's complete scuba and snorkeling store. The business offers underwater...Learn More
Tenkiller State Park is known as Oklahoma's "heaven in the hills." The sparkling blue waters of Lake...Learn More
Pensacola Dam and Grand Lake O' the Cherokees provide flood control for the Grand River and produce hydroelectric power...Learn More
Surrounded on three sides by the gorgeous Ouachita National Forest, Copper Sky Ranch is a unique getaway...Learn More
Dive into the clear, deep lake waters of Broken Bow Lake with the help of Broken Bow Scuba. Open by appointment...Learn More
Extreme Tornado Tours, based in Norman, has been featured on the Discovery Channel's TV show "Storm Chasers" for their...Learn More
This recreation area, covering portions of LeFlore and McCurtain counties in southeast Oklahoma, offers more than...Learn More
Pull through the gates of the Rutt-N-Strutt River Ranch near Lamar and check out of the daily grind and in to...Learn More
Located south of Weatherford, Crowder Lake offers 158 surface acres of water for recreational use. This lake has...Learn More
Hogan's Off Road Park in Disney is an off-road campground and recreation area perfect for any ATV or ORV. Located...Learn More
A prime example of Oklahoma's diverse terrain, Little Sahara State Park boasts over 1,600 acres of sand dunes,...Learn More
Lost Lakes Amphitheater & Water Park is a unique water sports park where you can wakeboard, waterski or kneeboard...Learn More
The Lake Eufaula Damsite ATV Trail features 300 acres of Canadian River Valley sand, rolling dunes and wooded...Learn More
The facilities at Lake Frederick include boat ramps, a boat dock, fishing dock, RV and tent campsites, picnic areas,...Learn More
Robbers Cave State Park is located in the scenic, hilly woodlands of the Sans Bois Mountains of southeast Oklahoma. This...Learn More
Located just south of the downtown area and the Oklahoma River, this 26,000 sq ft skate park is the largest and most...Learn More


text:
Lucky Brand
Geo Tile Blouson Top (Plus Size)
Product not available
Have questions?
Chat with us or call 1.888.282.6060.
HideDetails & Care
A three-button split-V placket and peekaboo trim detail a supersoft jersey top patterned in illustrative geo tiles. For an easy and flattering fit, a drawstring side tie cinches the blouson style.
- 28 1/2" length (size 2X).
- 60% cotton, 40% modal.
- Machine wash cold, tumble dry low.
- By Lucky Brand Dungarees; imported.
- Encore.


text:
Views of Sanders Reef, Bowling Ball and Schooner Beaches and miles of crashing surf on bluffs. Over two acres of amazing setting. Design and build your coast home on one of the finest sites to come along in years. Beach access to the south and the north, hiking trail along the bluff. Just south of Point Arena for shopping, dining, boat launch. Area fishing and diving. Wonderful full time living or coastal getaway.
Listing provided by Sharon Burningham at Coldwell Banker Coastal Properties
This Listing is for Sale .The listing status is Active, considered Resale, and located within Mendocino County
Copyright 2015, Bay Area Real Estate Information Services, Inc. All Right Reserved


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Neighbour services.
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text:
Newegg.com - A great place to buy computers, computer parts, electronics, software, accessories, and DVDs online. With great prices, fast shipping, and top-rated customer service - once you know, you Newegg.
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This review is from: G.SKILL Ripjaws X Series 4GB (2 x 2GB) 240-Pin DDR3 SDRAM DDR3 1333 (PC3 10666) Desktop Memory Model F3-10666CL9D-4GBXL
Pros: seemed to work fine, for a while.
Cons: after a month one of the chips started giving bsd. was after standard newegg return & had to rma to gskill.
Other Thoughts: never had a problem with gskill before. the rma was handled very fast.READ FULL REVIEW


text:
OVERVIEW Persistent changes in gene expression may mediate many effects of alcohol including reward learning, tolerance and dependence, suggesting that agents effective in changing alcohol-induced gene expression could be considered as therapeutic agents. Drugs targeting gene expression through inhibition of enzymes that regulate chromatin structure (epigenetic drugs) have been widely used in cancer research and recently emerged as potential therapeutics for neurodegenerative disorders and drug addiction. The main goals of this project are: 1) to identify epigenetic drugs that affect alcohol reward through testing their effects on alcohol consumption and conditioned place preference (CPP) and 2) to investigate the effects of selected epigenetic drugs on gene expression and cellular physiology in the reward pathway including the ventral tegmental area (VTA) and the nucleus accumbens (NA). The overall hypothesis is that some epigenetic drugs will reduce the rewarding properties of alcohol through changes in gene expression and cellular physiology in the reward pathway. Integration of electrophysiological and gene expression data will elucidate the drug's mechanisms of action and identify novel targets for drug development. This research will provide initial mechanistic evidence for the therapeutic potential of epigenetic drugs in treating alcohol addiction.
|Most, D; Ferguson, L; Blednov, Y et al. (2015) The synaptoneurosome transcriptome: a model for profiling the emolecular effects of alcohol. Pharmacogenomics J 15:177-88
|Cornelison, Garrett L; Mihic, S John (2014) Contaminating levels of zinc found in commonly-used labware and buffers affect glycine receptor currents. Brain Res Bull 100:1-5
|Mayfield, Jody; Ferguson, Laura; Harris, R Adron (2013) Neuroimmune signaling: a key component of alcohol abuse. Curr Opin Neurobiol 23:513-20
|Kirson, Dean; Cornelison, Garrett L; Philpo, Ashley E et al. (2013) Physiological concentrations of zinc reduce taurine-activated GlyR responses to drugs of abuse. Neuropharmacology 75:286-94


text:
Universal Fence Contracting & Landscaping
contracting, Site Servicing
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Lionel Messi has responded to his reports in the Spanish press that he made a gesture after being substituted in Barcelona's 4-1 home victory over Real Sociedad, claiming that some portions of the media had been "telling lies".
The forward, who had scored in the win, was taken off with nine minutes remaining by head coach Gerardo Martino, and Madrid-based sports newspaper AS reported that Messi was angry with Martino for substituting him.
"Nobody likes to be replaced, but we must accept it because it is the best for our team," Messi wrote on his Facebook page after the match.
"What I will not accept is that certain media tell lies about any situation that happens around me."
Martino told reporters after the match: "Messi never wants to be subbed off, but it's my task to take care of him. We need him at end of season as well as now."
It was the first time that Messi and recent signing, Neymar, had both scored for Barca scored in the same game, after the £49m transfer from Santos opened the scoring in the rout at the Nou Camp.
Messi has carried on his impressive form into this season, scoring seven goals in La Liga as Barcelona sit top of the division on goal difference ahead of Atletico Madrid. Both clubs have won all six of their league matches this season.Reuse content


text:
Discovering new design is why we love a trade show. Earlier this month at Top Drawer, never seen before companies took centre stage. Textile designer Deryn Relph was one of these first-timers; here she tells her story to Annie Deakin.
I've been in business for just over a year so… was unsure how financially viable a trade show like Top Drawer would be for me. I design and produce knitted chairs, lampshades and cushions using UK sourced cotton and lambswool yarns. Charlotte Abrahams invited me to be part of 'Spotted', a select group of 12 exhibitors who have not exhibited at a trade show before. It was such a great chance to take my business a step further.
The highlight of the show was… meeting so many wonderful people, both visitors and potential customers.
The lowlight… was standing all day for three days. Despite comfy shoes, my feet were sore at the end of each day.
The stand that caught my eye was… Rice who supply bright, bold and colourful homewares.
Of the 'Spotted' exhibitors, I loved… 'All Lovely Stüff', especially their Donut coat hooks and Bear Face mirror. Their appeal is the underlying humour and simplicity of well-considered design.
I stumbled upon a stand… called Whitehide Ltd selling printed leather-bound notebooks and sketchbooks and stationery. Their products are added to my wish list.
I was showing my two collections… The newer collection 'Buttonbox' was inspired by my recent inheritance of my Nanna's button box. I have memories of playing with the buttons as a child, of the clothes Nanna wore, and the whole 'make do and mend' mentality certainly inspires my nostalgic approach. The older one 'Retro Rainbow' evolved from my final degree project and was inspired by 'the future potential of seeds' and the micro-photographic work of Rob Kesseler, in conjunction with Wolfgang Stuppy of The Millenium Seed Bank Project. My own colour palette developed from a personal study of seeds and fruit, and includes lime, orange, cerise pink, magenta, lavender, Bordeaux and mustard yellow.
The biggest challenge in setting up my business is… balancing the different roles from accounts and answering emails to styling photography and attending shows. Sometimes there's barely time to knit or design new things.
Working from home… makes it tricky to achieve a work-life balance. Self-employment is harder than anticipated.
My greatest success is… making commercially viable products without losing that individuality.
I'm most proud of… my re-invention of the chairs. I create fabrics and put them together in a way that converts something from a piece of discarded rubbish to a highly desirable, contemporary interior piece that is also still functional.
My style is… mix and mismatch with a definite retro feel. I grew up in the 70's. I find colour – and the power it has over our minds - fascinating.
Knitting appeals to me because… it allows me to create the fabric and item in one go, to my own design. I spent many frustrated shopping trips hunting for a fabric that I had in my head but could never find.
There is something magical about… turning a ball or cone of yarn into cloth. Hand knitting is incredibly relaxing and knitted fabrics have unique properties too. I exploit the various knit structures and the stretch to work around retro style furniture and unusual lampshade shapes.
The perception of knitting has changed… it has achieved an almost cult status.
I see a definite trend for… retro style, mid century modern, 1960's and 1970's. There's obviously a trend for 'home-made', for mixing old and new and for making more considered purchases with artisan pieces.
Interior trends are… slower than fashion with less seasonality. I keep an eye on colour and style details in fashion trends because they often filter through to interiors a year later. The colour orange, eclectic tribal style and funked up craft are good examples. Following the recent crochet prints on the catwalk, I'm sure crocheted interiors, with granny squares aplenty, will be a trend soon.
I admire… Ercol because their brand has re-emerged and regained popularity by tweaking older designs to make them very contemporary. I admire Le Corbusier and Eileen Gray and Arne Jacobsen. Their designs are still so incredibly relevant and contemporary, even nearly 100 years later in some cases.
The designer to look out for is… Liam Treanor. I met him at New Designers One Year On last year.
I love the work of… Ptolemy Mann. Her use of colour is phenomenal, and she's proved that textile design skills have relevance in the built environment too. Donna Wilson has achieved a lot by breaking into the world of interiors with knit, and also managing to keep respect as a designer.
Annie Deakin is interiors writer for the mydeco marketplace, an online shopping experience where you can search hundreds of home furnishings and accessories all in one place.
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About Bikaner
About Bikaner
Lying in the north of the desert state, the city is dotted with scores of sand dunes. Bikaner retains the medieval grandeur that permeates the city's lifestyle. More readily called the camel country, the city is distinguished.
Arts Crafts of Bikaner
The Rajasthani miniature evolved styles that became apparent in the different Kingdoms where it found patronage. Used initially as the illusration for the text, they evolved later as the portfolios of the life and times.
Founder of Bikaner
Rao Bikaji was the most lively and enterprising among the five sons of Rao Jodhaji of Marwar . Realizing that a struggle for his paternal inheritance was not likely to be fruitful, this adventurous price was determined.
History of Bikaner
Palace, Bikaner Travel & TourismCity, north-central Rajasthan state, northwestern India. It lies 240 miles (386 km) west of Delhi.The city was the capital of the former princely state of Bikaner. About 1465 Bikaner.


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Wego.com recently found 36 hotels in Natchez , United States. Book last minute hotel or find great hotel deals such as The Elms Bed and Breakfast for US$183 per night. Save more on cheap hotels with rates from as low as US$60 per night.
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The Mighty Thor is a woman. That may sound like a sexist insult roared by the Hulk during one of the Marvel superheroes' many seismic punch-ups, but it's actually a statement of fact.
Marvel has announced that the God of Thunder, as played by Chris Hemsworth in the film franchise, is due to become a Goddess of Thunder later this year. And he (or she) isn't the only comics icon to find that the times are a-changin'. Also this week, in what read like a mis-timed April Fool's joke, Archie, an all-American teenager since 1941, died in Life With Archie issue number 36 when he intercepted an assassin's bullet intended for a gay senator.
That's not all. Marvel has just rebooted Ms Marvel as a Pakistani-American Muslim, and it recently unveiled a superhero team featuring no white males. Are mainstream comics finally embracing political correctness? Or – just throwing the idea out there – are they merely making headline-friendly changes in a desperate bid for free publicity?
"You can be cynical about it," says Chris Thompson at London's Orbital Comics shop, "but, in reality, these changes are more about redressing the balance. Up until now, superheroes have been straight white males, so by introducing gay characters, varying their ages, and putting in more women, they're just reflecting the diversity of the readership. This way, anyone who walks into a comic shop can find a character they relate to."
As for Thor's gender swap, says Thompson, it's in keeping with the character's mythology. "The idea is that anyone who picks up Thor's hammer gets the power, as long as they're worthy of it. He's been a frog before, he's been an alien, so why not a woman? And actually they've been building towards this within the series for quite some time. It's a natural evolution rather than a sudden switch."
So not a sign of an industry on the skids, then? Thompson thinks not, and Danie Ware at London's Forbidden Planet megastore agrees. "Superhero comics are doing fantastically well at the moment," she says. "Sales are booming, and we put that down to the mainstream acceptance of geek culture in general. Superhero movies have been absolutely huge, too, especially the Marvel ones [Iron Man, Captain America, Thor, Avengers Assemble]. We find that the films draw people in, and then they want to explore all the platforms for those characters: they buy the comics, they collect the collectibles." And they attend, as thousands did in London last weekend, events such as the London Film and Comic Con 2014.
Take a glance at the sales figures, and Ware's positivity seems to be borne out. In America, comics made more than $500m (£292m) last year, which is over $100m more than they made in 2003 – even if it's a long way below the industry's 1993 peak of $800m. The current wave of superhero blockbusters seems to be taking comics up, up and away.
Well, maybe. But if you look at the sales of each individual title, comics still appear to be a niche commodity, even if the associated DVDs, T-shirts and lunchboxes aren't.
Marvel's best superheroes
Marvel's best superheroes
1/19 Black Widow
The Black Widow (Natasha Romanoff) played by Scarlett Johansson
2/19 Spider Man
British actor Andrew Garfield as Spider-Man
3/19 Captain America
Chris Evans in 'Captain America: The First Avenger'
4/19 Iron Man
Robert Downey Jr in Iron Man 3, who won favourite action movie star at the People's Choice Awards 2014
Iron Man 3
5/19 Wolverine
Hugh Jackman bears his claws and loses the plot in X-Men movie 'The Wolverine'
Twentieth Century Fox Film
6/19 Hulk
Edward Norton as Hulk in The Incredible Hulk
7/19 Thor
Chris Hemsworth as Thor. Recently Marvel revealed its plans to make the hammer-wielding Norse superhero Thor a woman
Rex Features
8/19 Hank
Nicholas Hoult as Kan in 'X-Men: Days of Future Past'
9/19 Raven Darkhölme
Jennifer Lawrence as Raven Darkhölme in X-Men: Days of Future Past
10/19 Hawkeye
Jeremy Renner as Hawkeye in 'Avengers Assemble'
11/19 Winter Soldier
Sebastian Stan as the villainous Winter Soldier in the 'Captain America sequel
12/19 The Avengers
The Avenger's superheroes
13/19 Cyclops
The optic blast hero has long been played by James Marsden
14/19 Professor X
Patrick Stewart landed the role as Professor Charles Xavier in Bryan Singer's comic-book blockbuster X-Men
Rex Features
15/19 Rocket Raccoon
Bradley Cooper voices Rocket Raccoon in Guardians of the Galaxy
16/19 Groot
Also known as the Monarch of Planet X, Groot is played by Fast and Furious star Vin Diesel in Guardians of the Galaxy
17/19 Drax
Known as 'the destroyer', Drax is played by actor Dave Batista in Guardians of the Galaxy
18/19 Gamora
A soldier and assassin played by Zoe Saldana in Guardians of the Galaxy
19/19 Guardians of the Galaxy
Bill Mantlo's comic creation on screen in Guardians of the Galaxy
"There's the odd superhero comic that breaks 100,000 copies," says Paul Gravett, the co-curator of the British Library's Comics Unmasked exhibition. "But most of them sell well below that. The movies and TV show are helping, of course, but superhero comics are really quite a minor part of the product range now. They're really only useful as a source of reboots which can then be made into movies and cartoons."
But surely even Gravett – a champion of indie graphic novels – would acknowledge that a female Thor is a step in a more progressive direction? "Have you seen the pictures?" he asks with a sigh. "Show a woman in the street that crap image of Thor with tits, and ask her if she would read something like that. Of course she wouldn't. It won't make a blind bit of difference."Reuse content


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Honda confirms production Civic Type-R at Geneva
Honda confirms production Civic Type-R at Geneva
Meet the fastest and most powerful front-engined Ferrari ever, the 812 Superfast
Report: Alfa Romeo cans plans for a Giulia Sport Wagon
The all-new, Nurburgring-tuned 2018 Hyundai Elantra GT arrives as America's i30
Nissan extends the 2018 Titan lineup with a King Cab configuration
The new Ford GT specs are out and it's officially the fastest Ford ever made
Help the Dodge Viper ACR retake its title as the fastest production car around the Nürburgring
The 2018 Ford Mustang, this is it, with a new face
The Volkswagen I.D. Buzz is yet another bus concept for Detroit
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Latest Reviews


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The environmental plans for Skåldö, Högsåra, Iniö and Vänö are now done. Download by clicking on the brochure.
The Finnish environmental plans are now ready to be spread on the Islands. Here are the four different plans in finnish for Vänö, Iniö, Skåldö and Högsåra. Download by clicking on the brochure. Read more…
Kajsa Mellbrand from Aronia Research and Development Institute have summarized the results from the mapping of the ecosystem services on the participating islands. Download the report here>>
The Green Islands project arranged an information meeting on Ingmarsö on the 30th of November.Karin Larsson from Österåkers municipality talked about different waste water treatment techniques and how the municipality works with private sewage treatment. Cecilia Wibjörn from the Archipelago foundation talked about the project results and what we have done on Ingmarsö. Read more…
The Green Islands project are coming to an end. The Archipelago foundation have put together information for the Islands,Ingmarsö, Utö and Ornö. The information is a summation of experience and what we have learned over the past 2,5 years. Read more…
The Green Islands project have been gathering information about waste water treatment techniques, garbage handling, recycling, eco-system services, energy techniques and much much more. The project are now coming to an end and we want to share with you what we have learned so far. Where: Lurkan on Ingmarsö When: 30 November 12-15 Coffee and Read more…
The Green Islands project have been gathering information about waste water treatment techniques, garbage handling, recycling, eco-system services, energy techniques and much much more. The project are now coming to an end and we want to share with you what we have learned so far. Here is the invitation (PDF in Swedish)>> Where: Societetshuset on Read more…
Sustainable Saaremaa and Association of Estonian Islands invited stakeholders to an open discussion seminar. The seminar was held on Muhu on the 22 of October. They talked about tourism, low carbon transportation, infrastructure and the human relationship to nature. Key words for the day was intensive agriculture, small industries and local products (handicraft and food) Read more…
The Archipelago foundation held Green Islands final seminar on Monday October 7th. It was a day filled with information and inspiration. The three focus areas energy, waste water and garbage disposal were covered by four speakers. The project leader Cecilia Wibjörn shared the Green Islands project results. Gustav Hemming, Chairman of the Archipelago foundation Read more…
Kajsa Mellbrand fom Aronia, Finland was invited to talk about ecosystem services on the International Association for Landscape Ecology (IALE) conference on the 5th of September.
About the project
Questionnaire for the Swedish Islands in Green Islands.
Här är kartläggningsenkäten för de svenska öarna i Green Islands. Enkäten är uppdelad i en faktadel och en intressedel. De ska bara fyllas i en gång per hushåll, eventuell verksamhet som jordbruk, marina, bageri eller liknande ska inte räknas in i hushållet.
- December 2013
- November 2013
- October 2013
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- December 2012
- November 2012
- October 2012
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- June 2012
- May 2012
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- February 2012
- January 2012
- December 2011
- November 2011
- October 2011
- September 2011
- August 2011
- July 2011
- June 2011


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Thursday, October 6, 2016
Wednesday, September 14, 2016
Thursday, September 1, 2016
Thursday, August 18, 2016
Lauzier F.H.B Trailer
Tuesday, May 24, 2016
Lauzier FHB Logo/Website
Sunday, March 13, 2016
Friday, February 26, 2016
Monday, February 15, 2016
21516 : Foreign Elements
Tuesday, December 15, 2015
:Custom Vinyl Stormtrooper Head :
Wednesday, November 18, 2015
Sunday, November 8, 2015
Dirty Heads_Burn SLow
|I submitted a design to the Dirty Heads contest. The winners design will be made into merchandise and a portion of proceeds from sales will go to Oxfam America, a charity that works to combat against hunger, poverty, and social injustice. I've never had the opportunity of having my work be involved in something much bigger, so If selected as winner, I'd be stoked to see that through art I was able to make at least a little difference in the world. Any votes would be greatly appreciated
Wednesday, October 7, 2015
MiNd DuMP 20:15
Tuesday, September 29, 2015
Promotional Edit for Resyn Wakeskate's 2016 board graphic release
Tuesday, September 22, 2015
Friday, September 11, 2015
Thursday, September 10, 2015
"The Co-Pilot"
Sea bags are made out of recycled sails from ships and hand crafted in the heart of Portland, Maine. I was asked by an old friend who made the move from Maine back to Florida to create an image of her best friend that was making the trip by her side in the new jeep, his name is Max. Man's best friend is always so full of life and character so using the refrences I kept the design simple for threading purposes and created an illustration to match the trip they were about to embark on.
Custom sea bags can be purchased at www.seabags.com
Monday, July 13, 2015
Captain Vs Chaos
Chaos "Dingley Sticker" of Captain Dingley's Board Shops mascot.
Captain Dingley's Board ShopNaples Original Watersports Location! 207-693-LAKE Find us on Facebook! No Shirt, No Shoes, No Worries!
Current Project With Resyn Wake
The past few weeks I've been bringing to life some really fun and awesome concepts that will be coming out for Resyn Wakeskates new line in the coming months. Its been an awesome adventure full of learning and pushing my style while working with color a lot more and keeping a very iconic feel with a dab of graffiti influence. Keep and eye out on Resyns Instagram and website so you don't miss out. These are going to be killer!
Saturday, April 25, 2015
"Hello My Name Is ?" 2015
Friday, April 17, 2015
Wednesday, April 8, 2015
Sunday, February 15, 2015
Crypt Keeper
Monday, May 6, 2013
Wednesday, February 27, 2013
Thursday, February 23, 2012
Absence Of The Sun Logo
Monday, January 2, 2012
Esoteric Burial T Design
Sunday, July 3, 2011
Monday, May 16, 2011
Snowboard Mock Ups
Friday, March 11, 2011
Tuesday, October 26, 2010
Monday, April 19, 2010
Brzowski Logo
|concept sketch
|Hand Drawn Logo in Photoshop
Final logo as seen on cd release
by Brzowski
20 tracks, 71 minutes
released 01 November 2012
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Another dry winter could hurt Arizona's water supply
PHOENIX — Arizona is facing the possibility of a third consecutive dry winter — that's not good news for the state's water supply.
Mark Hubble with SRP said the reservoirs that supply the Valley with water are below 50 percent of capacity. The snowpack in Colorado, Utah and Wyoming is half of what it should be. That snowpack runoff feeds the Colorado River, which Arizona taps into via the Central Arizona Project canal.
"[That is] A concern for us because the Colorado Basin really supplies the entire southwest. Phoenix is fortunate that we have our own watershed." But the runoff is still needed.
Another dry winter could tax the water supply starting late next summer but it's too soon to start discussing cutbacks and possible rationings.
"If we have another dry winter again I think you'll start hearing some rumblings about the surface water supply," Hubble said.
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- Operation Santa Claus needs holiday help
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- Arizona kids in crisis: How you can help
- 11 holiday classics for the ultimate movie marathon
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Government mutual will be a beacon for IT procurement
The pension administration mutual being set up by the government will be the first of a wave of new business models that could shake-up central government outsourcing contracts.
This Article Covers
Government & public sector IT
The pension administration mutual being set up by the government will be the first of a wave of new business models that could shake-up central government outsourcing contracts.
EU data protection: Essential guide
Access this guide to aid you in facing the challenges, opportunities and key actions need to consider in preparing your organisation for GDPR.
By submitting your personal information, you agree that TechTarget and its partners may contact you regarding relevant content, products and special offers.
You also agree that your personal information may be transferred and processed in the United States, and that you have read and agree to the Terms of Use and the Privacy Policy.
Speaking at The Crown and suppliers: A new way of working, a government procurement conference in London, Katharine Davidson, director of strategy in the Cabinet Office Efficiency and Reform Group (ERG), said the My Civil service Pension (MyCSP) mutual currently being set up will be the first example of a new way the government will work with suppliers.
MyCSP was announced earlier this year as the first example of a government body spun out into a mutual and partnered with a private sector company. MyCSP will administer 1.5 million civil servant pensions, and will be owned by three groups: the 475 staff, the government and a private company that will run the service. Four companies were on a shortlist to be the private sector partner that would run the scheme but Indian IT services Wipro pulled out leaving Capita, Xafinity and JLT to fight it out.
Davidson said mutual and joint ventures, which retain a government stake, will be more common in the future as the government strives for "better" service arrangements.
"Everything is up for grabs and the pie is bigger," she told an audience of 800 representatives from suppliers.
She said the value in mutuals, which she says makes them a better option, is driven by the fact that the success of the organisation is in the interest of all the stakeholders: taxpayers, staff and the private sector partner.
Taxpayers will get better value and post-deal services, according to Davidson. Meanwhile employees will have an ownership stake and as a result will be more likely to innovate, and the private sector partner will be able to engage more with the public sector.
She said this will help increase public sector productivity levels: "The new business models will help the public sector close the productivity gap [that exists between it and the private sector]."
The government will announce a new position early next year, known as head of commercial at ERG, to drive the strategy, which will create a "pipeline of opportunities for new models," said Davidson.
Enjoy the benefits of CW+ membership, learn more and join.
European law enforcement seeking smart ways to fight cyber crime
Read more on IT for government and public sector
Gov.uk Verify ID providers eye the prize of HMRC's digital users – but what happens without them?
HMRC ID vs Gov.uk Verify – what's the difference, and why it matters
GDS, HMRC and Verify: so much for cross-government digital collaboration
The government's transformation strategy is ambitious, but is it achievable?


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More Videos...
Hot Rod and Stock Eliminator Highlights @ IHRA North American Nationals
IHRA North American Nationals at New England Dragway in Epping NH. Even though it was rain shortened on Friday Sept. 11, 2009, there were some awesome runs in all classes. Here are some nice cars and wheels up run from Hot Rod (10.90 index) and Stock Eliminator.
Super Stock IHRA Nitro Jam New England Dragway 2010
First Round Qualifying and First Round Eliminations from The IHRA North American Nitro Jam Event at New England Dragway in Epping, NH Sept. 11, 2010
2011 NHRA AAA Auto Club Finals Stock Eliminator 1st Round
Entire 1st Round of Eliminations
Stock Eliminator Round 1 Eliminations LODRS Maple Grove 2012
Round one qualifying at the Lucas Oil Drag Racing Series Event at Maple Grove Raceway in PA.Saturday May 26, 2012. My apologies, I found out what my new camcorder doesn't like...as I sat through about five pairs of cars and didn't have access to record while it was restoring files...and naturally two of the best Stock wheelstands of the weekend occurred as I watched and couldn't do anything about it!


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Exclusive, Timeless Affairs
We invite you to celebrate your special moment in the sophistication of Vdara Hotel & Spa. When you host your private wedding reception, anniversary, or glamorous social affair at Vdara, the space and experience is intimately yours. With personal attention to every detail, enjoy all-suite accommodations to host and prepare for your special day. Vdara's unique venues include sophisticated ballrooms or an intimate pool deck with spectacular city views, exclusive access to the Spa & Salon, limousine service and more. Toast your unique moment, dine and dance in a glamorous escape, only steps away the center of excitement in Las Vegas.
Catering & Banquet Sales 702.862.1559
General Inquiry 866.745.0262
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Add your personal touch with flowers and musical accompaniment.
Contact us at 866.745.0262 or 702.590.2325
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Soothe and transform in the peaceful
ESPA at Vdara & Salon Vdara.
To schedule an appointment, call 866.745.5654 or 702.590.2474
The Wedding Chapel at ARIA
Enjoy the luxurious amenities of ARIA as you and your special someone say 'I Do'. Explore all The Wedding Chapel has to offer for your special day.
ARIA Weddings
Weddings/Social Occasions Brochure
Special Accommodations
Escape to your stunning hospitality suite to bask in your moment, choose from several suite plans, room blocks or any special arrangement you desire.
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Successful Meetings at Vdara
Meetings held in comfort and convenience.
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Meetings at Vdara
Vdara's exceptional and intimate service extends to the meeting spaces – specializing in smaller groups from 10 to 300 guests.
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Vdara features a variety of ideal settings including our spacious Hospitality Suites, the whimsical VICE VERSA Patio & Lounge, and Pool & Lounge Vdara featuring cabanas, private retreats and stunning views.
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Updated policies and procedures for your meetings at Vdara.


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Jovinna Chan, E-RYT 500,
is a life mentor and co-founder of Shakti Initiation, and leads transformational programs around the world. She is a faculty member of Pranotthan School of Yoga, served as the assistant dean, 200- and 500-hour teacher trainer for the Kripalu School of Yoga, and she's also certified in LetYourYogaDance and Soul Motion™.
Legacy Group - Coming Alive through Conscious Dance, Meditation and Yoga Philosophy
Are you ready to tap into your potential for deeper love, awareness, aliveness, and creativity? Then pause from your routine, and take on the role of a devoted apprentice to mastering the mystery and science of being YOU.
Fully Alive: Unveil Yourself through Mindfulness, Yoga and Conscious Dance
Who am I? What is my purpose? What is the most importing thing right now in my life? Have you asked yourself these questions? According to the Bhagavad Gita, an ancient yogic scripture, it is better to live your own dharma (calling) imperfectly, than to live someone else's dharma perfectly. Sometimes, however, the greatest challenge is recognizing your dharma. The Gita and your own body offer guidance.