Advances on antibiotic resistance genes (ARGs) in aquaculture environment
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摘要: 抗生素对水产养殖业中水生生物疾病防治、生产线增产等发挥着重要作用，但长期滥用抗生素很可能会诱导水生生物体内产生携带抗生素抗性基因 (Antibiotic resistant genes, ARGs) 的耐药菌 (Antibiotic resistant bacteria, ARB)。ARGs在水产养殖环境中的持久性残留、迁移和传播，会埋下基因污染隐患，导致生态失衡并危害人类安全，如何遏制抗生素抗性的传播已引起全球重点关注。就水产养殖环境中ARGs的研究进展，系统总结了ARGs的污染现状及其在水产养殖环境中的来源、迁移传播和影响因素，并简述了ARGs与抗生素、微生物群落和环境因素之间的关联特性，以及抗生素、ARGs和ARB对生态环境与人类健康的影响。基于此，概述了ARGs的控制策略与去除技术，并提出了今后的研究方向，以期为水产养殖环境中ARGs污染机理的解析和抗生素抗性传播风险的控制提供科学参考。Abstract: Antibiotics play a significant role in the disease control of aquatic organisms and output increase of aquatic products. However, long-term abuse of antibiotics can result in the occurrence of antibiotic resistant bacteria (ARB) which harbor antibiotics resistance genes (ARGs) in aquatic organisms. The persistent existence, migration and spread of ARGs in aquaculture environment will potentially cause genetic pollution, destroy the ecological balance, and pose risks to human health. Therefore, how to constrain the spread of antibiotic resistance has attracted global attention. In terms of the research advancement of ARGs in aquaculture environment, this review systematically summarizes the status of ARGs pollution coupled with the source, migration and spread behavior of ARGs and their influencing factors, illustrates the correlations between ARGs and antibiotics, microbial communities and environmental factors, as well as discusses the effect of antibiotics, ARGs and ARB on ecological environment and human health. Thus, the paper reviews the management strategies and removal technologies of ARGs, and proposes the future research directions regarding ARGs, so as to provide references for revealing the pollution mechanism of ARGs and reducing the transmission risk of antibiotic resistance.
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黄唇鱼(Bahaba taipingensis)属石首鱼科，为近海大型暖温性底层鱼类，分布于中国南海和东海，尤其在珠江口较为常见，是中国特有种，1988年被列为国家二级重点保护水生野生动物[1-2]。黄唇鱼鱼鳔(俗称“鱼胶”)被认为有特殊的药用价值，自古以来一直是药用及滋补极品，极为珍贵[3]。黄唇鱼曾是珠江口重要的渔业捕捞对象[4]，但20世纪后期以来，由于受到该海域水环境污染、高强度开发和非法捕捞等的影响，其资源量急剧下降，现处于濒临灭绝状态，2006年被世界自然保护联盟(IUCN)物种红色名录列为极度濒危物种(CR)[5]。
中国关于鱼类发声的研究在20世纪80年代已有开展[6-8]，但其后一直发展缓慢，鲜有相关研究报道；且国外多研究鲸豚类发声，对鱼类发声特性的研究也较少报道[9-14]。石首鱼科鱼类能依靠鳔的振动发出明显的声音[15]，其发声行为与生殖、防御、索饵等活动有密切关系[16-19]，因而为研究者所关注。黄唇鱼和大黄鱼(Larimichthys crocea)均属石首鱼科，近年来已有使用被动声学方法研究大黄鱼、褐菖鲉(Sebastiscus marmoratus) 等的发声特性，但由于黄唇鱼的稀有性，目前尚未见对其发声特性的研究报道。
声学探测包含被动探测与主动探测。使用被动声学方法探测黄唇鱼发声特性，既不会对鱼造成伤害，也不会破坏海洋环境。因此，本文通过被动声学方法监听黄唇鱼的声音，初步分析了其声谱特征，旨在为黄唇鱼声学无损调查、水下噪声影响分析和发声的生物学行为等物种保护研究提供基础数据。
1. 材料与方法
1.1 实验仪器与实验方法
实验仪器为microMARS水听器，前置放大增益为18 dB，平坦频率响应范围为0.6~33 kHz，信号范围为70~166 dB re 1 μPa，用电池供电。
2017年3—5月，对东莞黄唇鱼市级自然保护区救护驯养基地的室内水族箱和室外驯养池中黄唇鱼的水下发声情况进行了水下监听。室内水族箱高215 cm、长420 cm、宽150 cm、水深2 m，其中放养2尾体质量约25 kg的黄唇鱼，室内自然光，夜间不开灯。室外驯养池为矩形池塘，水深2~2.5 m，1#池面积约3 000 m2，驯养有29尾体质量约15~25 kg的黄唇鱼；2#池面积约6 300 m2，驯养有65尾体质量约8~13 kg的黄唇鱼。水听器固定在保护架上，放在水族箱底部一角，驯养池放置在池中央水深约2.5 m的池底。水听器采样频率为30 kHz，水族箱和2个训养池均监听2次，每次7 d。每次回收水听器后，导出并保存监听时间段内的WAV音频文件。
1.2 音频数据分析方法
通过音频软件Cool Edit Pro 2.1和Audacity试听音频并观察波形图，确定声信号类型，并对不同类型声信号、波形和时长等参数进行统计分析。白昼、夜晚的起始和结束时间以日出和日落时间为界线[20]。用MATLAB R2012a软件对声音信号进行时频分析，画出声音信号波形图；对声音信号进行短时傅立叶分析得到语谱图，亦称为可视语音；对声音信号作傅立叶变换得到频谱图，其中频谱图中的峰值对应的频率为谱峰中心频率[21-23]。
2. 结果
2.1 发声类型
共获得音频文件总时长907.54 h，通过音频文件试听，结合声音信号的波形变化比对，将黄唇鱼发声分为7类：类鼓声、咔嚓声、雀鸣声、嗡嗡声、嗒嗒声、嚓咕声和其他声(图1)。监听期间共监听到黄唇鱼发声246次，其中类鼓声175次，约占71.1%，咔嚓声占比6.9%，雀鸣声、嗡嗡声、嗒嗒声、嚓咕声等占比均不超过4%，其他声音占比15.4% (表1)。其他声音主要出现在第二次水族箱监听中，共26次，且大多为不同声音。室外训养池与室内水族箱中黄唇鱼平均发声密度变化范围为0.040~0.712次·h–1，均随时间呈增加趋势(表1)。黄唇鱼中发声次数成对样品t检验分析表明，1#、2#池和水族箱中黄唇鱼的发声次数均无明显差异(P>0.18)。类鼓声包含的脉冲数变化范围为1~3个(图2)，以单脉冲为主(139次)，约占类鼓声总数的79%，类鼓声为类正弦波形，单脉冲类鼓声含3~43个波形；其他类别声音为单脉冲或多脉冲声音。
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图 1 黄唇鱼主要发出的6类声音a. 类鼓声；b. 嗡嗡声；c. 咔嚓声；d. 嗒嗒声；e. 雀鸣声；f. 嚓咕声Figure 1. Six kinds of sounds made mainly by B. taipingensisa. drum sound; b. humming sound; c. cracking sound; d. clacking sound; e. bird sound; f. cha goo sound![]()
图 2 单脉冲 (a)、双脉冲 (b) 和三脉冲 (c) 类鼓声Figure 2. Single pulse (a), double-pluse drum sound (b) and three-pluse (c) drum sound表 1 各类别发声次数Table 1. Number of different sound types监听区
monitored area时间 (月-日)
time (month-date)总发声数
total number of sounds
平均发声密度/次·h–1
mean sound density类鼓声
drum sound咔嚓声
cracking sound雀鸣声
birds sound嗡嗡声
humming sound嗒嗒声
clacking sound嚓咕声
cha goo sound其他声
other sound1#池
Pool 103-28—04-04 7
0.0400 0 0 6 0 0 1 04-21—04-28 10
0.0587 3 0 0 0 0 0 2#池
Pool 204-05—04-12 19
0.1115 6 1 0 5 0 2 05-22—05-29 63
0.36450 5 0 2 0 0 6 水族箱aquarium 05-02—05-04 24
0.52221 0 0 0 0 0 3 05-10—05-17 123
0.71292 3 1 0 0 1 26 合计 total 246 175 17 2 8 5 1 38 2.2 昼夜发声比较
实验监听期间，白昼监听到黄唇鱼发声134次，夜晚监听到112次，其中白昼类鼓声96次、夜晚79次。5月前监听夜晚发声次数多于白昼，之后白昼发声次数多于夜晚(图3)，但成对样品t检验分析表明，黄唇鱼昼夜发声次数并没有明显差异(P=0.12)。
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图 3 总发声和类鼓声昼夜分布Figure 3. Day and night signal distribution of total sound and drum sound2.3 类鼓声声谱特征
2.3.1 类鼓声时频特征
黄唇鱼的类鼓声语谱图显示其声信号能量集中在0~1 000 Hz，其声纹与时间轴平行(图4)；类鼓声频谱图显示其谱峰中心频率集中在50~140 Hz (图5)。
2.3.2 类鼓声时域特征
黄唇鱼的类鼓声时长范围为67~1 333 ms，总平均值为279 ms，众值为100 ms，67~533 ms时长段类鼓声占93% (图6)；类鼓声的时长分布符合等差数列公式t=33.3+33.3n [t为时长(ms)，n为正整数]。类鼓声脉冲宽度和脉冲间隔范围分别为35~733 ms和0~1 130 ms，总平均值分别为70 ms和183 ms。单脉冲类鼓声时长(脉冲宽度)范围为67~733 ms，平均值为243 ms；双脉冲类鼓声时长范围为100~1 333 ms，平均值为370 ms，脉冲宽度范围为35~350 ms，平均值为113 ms，脉冲间隔范围为0~1 130 ms，平均值为143 ms；三脉冲类鼓声时长范围为333~1 268 ms，平均值为655 ms，脉冲宽度范围为47~333 ms，平均值为100 ms，脉冲间隔范围为0~834 ms，平均值为179 ms (表2)。
表 2 类鼓声时域特征统计Table 2. Pulse width and pulse interval of drum sound脉冲类型
number of pulse时长/ms
duration time脉冲宽度
pulse width脉冲间隔
pulse interval单脉冲 single pulse drum sound 67~733 243 67~733 243 − 双脉冲 double-pulse drum sound 100~1 333 370 35~350 113 0~1 130 143 三脉冲 three-pulse drum sound 333~1 268 655 47~333 100 0~834 179 总平均值 overall mean 279 70 183 2.3.3 其他发声类别时频特征
黄唇鱼发出的类鼓声、咔嚓声、雀鸣声、嗡嗡声、嗒嗒声、嚓咕声等6类声音的语谱图及频谱图见图7。嗒嗒声的语谱图与类鼓声相似，显示其能量也集中在0~1 000 Hz，声纹与时间轴平行(图7-d1)，但其谱峰中心频率范围为180~190 Hz (图7-d2)。嗡嗡声、咔嚓声、雀鸣声、嚓咕声能量和频率分布均范围较广，在低频和高频均有分布；其中嗡嗡声能量集中在0~1 000 Hz、2 000~6 000 Hz (图7-b1)，谱峰中心频率范围为40~140 Hz (图7-b2)；咔嚓声能量集中在2 000~5 000 Hz (图7-c1)，谱峰中心频率范围为3 200~3 600 Hz (图7-c2)；雀鸣声能量集中在0~3 000 Hz、10 000~12 500 Hz (图7-e1)，谱峰中心频率有2处，范围分别为400~500 Hz和2 000~2 500 Hz；嚓咕声能量集中在0~5 000 Hz(图7-f1)，谱峰中心频率范围为50~150 Hz (图7-f2)。
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图 7 语谱图和频谱图a1~a2. 类鼓声语谱图和声频谱图; b1~b2. 嗡嗡声语谱图和频谱图; c1~c2. 咔嚓声语谱图和频谱图; d1~d2. 嗒嗒声语谱图和频谱图; e1~e2. 雀鸣声语谱图和频谱图; f1~f2. 嚓咕声语谱图和频谱图Figure 7. Spectrogram and spectrum mapa1−a2. spectrogram and spectrum map of drum sound; b1−b2. spectrogram and spectrum map of humming sound; c1−c2. spectrogram and spectrum map of cracking sound; d1−d2. spectrogram and spectrum map of clacking sound; e1−e2. spectrogram and spectrum map of bird sound; f1−f2. spectrogram and spectrum map of cha goo sound3. 讨论
3.1 发声机制
鱼类发声行为的机制分别为鳔发声、摩擦发声、呼吸发声或其他[15]。石首鱼科鱼类主要依靠气鳔的振动发声，它们都有两束来自腹腔并直接或间接与鳔相联系的肌肉，肌肉急速收缩和放松，就能使鳔振动发声，鳔还可以作为共振器来帮助发声。鳔发声的强度和频率也可以调节。这种发声机制具有非常宽的频带，一般为几百至十几千赫兹。根据Sprague[24]的理论：发声鱼类的鱼鳔肌可视为振动弹簧模型，弹性肌肉的收缩和释放则分别对应于正弦波的不同半周期，而鱼鳔则作为一个阻尼结构，使鱼鳔肌的收缩逐渐衰减[25]。本研究中黄唇鱼的类鼓声脉冲波形图具有若干个类正弦波形，可以推测黄唇鱼类鼓声的发音类型属于鳔发声。此外，黄唇鱼类鼓声语谱图声纹与时间轴平行，显示了共振峰特征，根据语谱图声纹和共振发声的关系[26]，可以推测黄唇鱼主要是以鳔为共振器来帮助发声[15]。
3.2 发声类型
通常，不同种类的鱼所发出的声音在频率上差异较大，即使同一种类，由于其本身生物学状态的不同以及所处环境条件的变化，它们所发出的声音也各不相同[15]。珠江河口水域的生物发声监测辨认出66种声音类型，并倾向于拥有一个脉冲串结构[27]。大黄鱼觅食时发出的“咕噜噜”声信号都是简单的单脉冲，产卵时发出的“咯咯咯”声信号则大部分是连续的双脉冲或三脉冲，只有极个别为单脉冲或多脉冲[23]。大黄鱼也发出惊扰声和摄食声[28]。褐菖鲉在领地入侵实验中发出“咕噜噜”的叫声，其声音波形由几个单独脉冲或一组连续脉冲构成，波形特征基本相似[25]。与大黄鱼、褐菖鲉相似，本研究中黄唇鱼发出的声信号主要为单脉冲信号，也有双脉冲或三脉冲信号，主要发出类鼓声、咔嚓声、雀鸣声、嗡嗡声、嗒嗒声、嚓咕声等6类声音，类鼓声占比高达71.1%，由此可以推测类鼓声是黄唇鱼发声行为中最重要的类别，对种群内的信息传递有重要意义。
3.3 声谱特征
不同水生生物的声谱特征存在较明显的差异。鱼类发声频率一般为低频，谱峰中心频率多在1 000 Hz以内(表2)，珠江河口水域生物的66种声信号的谱峰中心频率范围为500~2 600 Hz，能量集中在4 000 Hz以下[27]。与鱼类不同，鲸豚类一般发出宽带高频声信号，如江豚(Neophocaena phocaenoides)发出的探测声信号谱峰中心频率为87~145 kHz[34]；中华白海豚(Sousa chinensis)发出的click串声信号谱峰中心频率为70~80 kHz，最高频信号甚至在125 kHz以上，主要能量分布在50 kHz以上[8,35]。黄唇鱼发出的类鼓声能量集中在0~1 000 Hz，谱峰中心频率为50~140 Hz，发出的嗒嗒声的能量分布及谱峰中心频率范围与类鼓声相似，发出的嗡嗡声、咔嚓声、雀鸣声、嚓咕声能量同时包含低频(0~1 000 Hz)与高频成分(3 000~12 500 Hz)，其谱峰中心频率范围也较广。与黄唇鱼同属石首鱼科的大黄鱼也会发出多种声音，如咕噜噜摄食声、咯咯咯产卵声和惊扰发声，其谱峰中心频率范围在630~800 Hz，梅童鱼(Collichthys lucidus)、黄姑鱼(Nibea albiflora)、白姑鱼(Argyrosomus argentatus)、叫姑鱼(Johnius belengeri)和拟石首鱼(Sciaenops ocellatus)等其他石首鱼科鱼类声信号的谱峰中心频率范围为139~2 000 Hz，非石首鱼科鱼类声信号的谱峰中心频率范围为83~1 200 Hz (表3)。可见，相比其他鱼类的声信号，黄唇鱼的类鼓声、嗒嗒声、嗡嗡声和嚓咕声的谱峰中心频率处于偏低水平。
表 3 部分鱼类声信号谱峰中心频率Table 3. Spectral frequency peak of sound signal of servaral fishes种类
species谱峰中心频率/Hz
spectral frequency peak文献来源
Reference黄唇鱼 Bahaba taipingensis 类鼓声50~140；嗒嗒声180~190；嗡嗡声40~140；咔嚓声3 200~3 600；
雀鸣声400~500、2 000~2 500；嚓咕声50~150本研究 大黄鱼 Larimichthys crocea 630~800 [7,23,28] 梅童鱼 Collichthys lucidus 1 000 [8] 黄姑鱼 Nibea albiflora 650±20.12 [23] 尖头黄姑鱼 Nibea acuta 630±15.57 [23] 白姑鱼 Argyrosomus argentatus 400 [29] 叫姑鱼 Johnius belengeri 2 000 [29] 拟石首鱼 Sciaenops ocellatus 139 [30] 金尾贝氏石首鱼 Bairdiella chrysoura 1 046 [30] 狗䱛 Cynoscion regails 347 [30] 云斑狗䱛 Cynoscion nebulosus 300 [30] 红牙䱛 Otolithes ruber 632±10.06 [23] 褐菖鲉 Sebasticus marmoratus 83~174 [25] 金眼鲷 Beryx splendens Lowe 337±8.50 [23] 带鱼 Trichiurus haumela 628±11.40 [23] 白鱼 Salangichthysm icrodon Bleeker 536±10.39 [23] 刺鱼 Gasterosteus aculeatus Linnaeus 420±0 [23] 大斑石鲈 Pomadasysmaculatus 415±9.40 [23] 粗纹鲾鱼 Leiognathus lineolatus 757±24.70 [23] 黑鳍叶鲹 Atule malam 542±16.90 [23] 游鳍叶鲹 Atule mate Cuvier et Va lenciennes 528±9.67 [23] 白舌尾甲鲹 Uraspis helvola 534±17.92 [23] 海鲶 Arius sp. 735±12.39 [23] 东方豹鲂鮄鱼 Dactyloptena orientalis 348±0 [23] 鳓鱼 Ilisha elongata 251±18.41 [23] 白腹豆娘鱼 Abudefduf luridus 356 [31] 鼬鳚 Ophidion marginatum 1 200 [32] 红棘胸鲷 Gadidae mediterraneus 180 [33] 黄唇鱼的声谱特征与其他鱼类相比，差异与共性并存，鱼类发声频率范围一般为低频(1 000 Hz以内)，但这并不意味着不同鱼的声谱特征、时域特征是重复的，如不同声音听觉上就完全不同，其波形、谱峰中心频率、语谱图等也均有明显差异，可根据这些差异进行鱼类发声分类，及其声谱和时域特征辨别。待鱼类声信号特征提取达到一定水平和累积足够多的鱼类声信号数据后，才可能对鱼类声信号表征的鱼类信息交流和行为进行精准鉴别。
3.4 时域特征
黄唇鱼发出的类鼓声时长范围为67~1 333 ms，总平均值为279 ms，众值为100 ms，时长分布符合等差数列公式t=33.3+33.3n [t为时长(ms)，n为正整数]；类鼓声脉冲宽度和脉冲间隔范围分别为35~733 ms和0~1 130 ms，总平均值分别为70 ms和183 ms。黄唇鱼类鼓声时长和脉冲间隔随着声信号脉冲数的增加而增长，脉冲宽度则减短。大黄鱼摄食声或产卵声的声信号时长1~2 ms，摄食声的脉冲间隔为1~30 ms，产卵声的脉冲间隔为90~140 ms，其时长和脉冲间隔均远比黄唇鱼类鼓声的短[23]。褐菖鲉平均脉冲宽度(32.6±2.6) ms[25]，也明显短于黄唇鱼类鼓声的平均脉宽。与鲸豚类比较，中华白海豚click声信号脉冲间隔变化范围为3.3~349.2 ms，也短于黄唇鱼类鼓声的脉冲间隔[35]。可见水生生物在发出的声信号时长、脉冲宽度和脉冲间隔等的差异主要与不同发声生物及不同声信号类型有关。
4. 结论
本文对人工圈养的黄唇鱼在不同实验条件下(室内水族箱、室外池塘)的发声信号进行采集，对黄唇鱼的发声机制、发声类型、声谱特征及时域特征进行了初步分析。结果表明，黄唇鱼声学监听共监听到246次发声，发出的声音分为7类，分别是类鼓声、咔嚓声、雀鸣声、嗡嗡声、嗒嗒声、嚓咕声和其他声音等。黄唇鱼昼夜发声次数没有明显差异。黄唇鱼发声以类鼓声为主(175次)，类鼓声由1~3个脉冲组成，又以单脉冲类鼓声为主(139次)。类鼓声为类正弦波形，能量集中在0~1 000 Hz，声纹与时间轴平行，谱峰中心频率为50~140 Hz。嗒嗒声与类鼓声相似，其能量也集中在0~1 000 Hz，但谱峰中心频率范围为180~190 Hz。嗡嗡声、咔嚓声、雀鸣声和嚓咕声同时包含低频(0~1 000 Hz)与高频(3 000~12 500 Hz)成分。类鼓声时长、脉冲宽度和脉冲间隔范围分别为67~1 333 ms、35~733 ms和0~1 130 ms，平均值分别为279 ms、70 ms和183 ms；类鼓声时长和脉冲间隔随着声信号脉冲数的增加而增长，脉冲宽度则减短。
实验用水听器平坦频率响应范围为0.6~33 kHz，而监听到低于600 Hz的黄唇鱼发声信号，说明仅距离水听器较近的黄唇鱼发声信号被监听到，距离水听器较远的黄唇鱼发声信号未被监听到。所以有必要对实验监听系统作改进，以监听并记录到更多的黄唇鱼发声信号和获得更全面的黄唇鱼发声特征。另外，不同实验条件下黄唇鱼发声信号可能存在一定差异。目前声纹识别技术研究仅限于人类语音学，还未应用在鱼类声信号研究中。在今后的工作中需要通过更多实验获取更加丰富的数据和改进数据分析方法进行深入研究，并结合黄唇鱼的行为特征来归类划分、建立数据库，为人工救护、喂养、保护珍稀黄唇鱼提供参考。
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图 1 水产环境中ARGs的来源、迁移与传播
Figure 1. Source, migration and spread behavior of ARGs in aquaculture environment
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图 2 水产养殖环境中ARGs与抗生素、微生物群落和环境因素之间的关联特性
Figure 2. Correlations between ARGs and antibiotics, microbial communities and environmental factors in aquaculture environment
表 1 现有技术对ARGs的去除效果
Table 1 Reduction efficiency of ARGs by existing technologies
去除技术
Removal technology去除原理
Removal principle去除效果
Reduction efficiency参考文献
Reference添加大孔吸附树脂
Adding macroporous adsorption resin (MAR)MAR是一种多孔交联聚合物，能够降低ARGs和微生物群落的丰度，并且通过吸附重金属以降低其对ARGs的协同效应和选择压力。 ARGs (14.14%~99.44%)和MGEs (47.83%~99.48%)的丰度显著降低。 [101] UV/氯消毒
UV/chlorineUV/氯协同作用可以有效灭活ARB、打破ARGs结构并抑制其水平转移。 UV (320 mJ·cm−2)/氯(2 mg·L−1)协同作用下，ARGs的去除率增强了1~1.5 log。 [102] 臭氧后处理
Ozone post-treatment臭氧具有高氧化电位 (2.07 V)，可以有效去除ARGs和ARB。 胞内ARGs (iARGs)的去除率达到89%。 [103] 高铁酸盐
Ferrate高铁酸盐作为一种高价铁基氧化剂，其强氧化电位能够直接去除ARGs，且具备较强的杀菌效能，能够灭活携带ARGs的细菌，从而抑制其垂直转移。 高铁酸盐的剂量为10 mg-Fe·L−1时，ARGs的去除率达到1.10~4.37 log。 [104] 生物过滤
Biofiltration水体中的微生物会附着在过滤介质 (石英砂、颗粒活性炭和无烟煤等) 表面并形成生物膜。 ARGs平均丰度降低了0.97 log。 [105] 污泥处理湿地
Sludge treatment wetlands (STWs)STWs法是传统沙干化床和垂直流人工湿地的联合技术，剩余污泥进入湿地后会形成不同污泥层，而植物在其中生长，有利于稳定污泥、减少污泥体积并去除ARGs等污染物。 磺胺类ARGs的丰度降低了21%。 [106] 
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