来源:Alexander S. Gold1,Tonya M. Colpitts1
1 微生物学系,波士顿大学医学院,国家新发传染病实验室,马萨诸塞州波士顿
转导是一种通过噬菌体介导的细菌间基因交换形式,噬菌体(或称噬菌体)是一类专门感染原核生物的病毒。这种通过噬菌体将DNA从一个细菌转移到另一个细菌…
1. 设置
2. 实验方案
3. 数据分析与结果
| 变性 | 时间 | |
| 变性 | 94 °C | 2 min |
| 40 个循环: | ||
| 变性 | 94 °C | 15 sec |
| 退火、延伸及荧光读取 | 60 °C 或比最低引物 Tm 低 5 °C | 1 min |
表1:qPCR热循环程序
细菌可以通过交换遗传物质快速适应快速变化的环境,其中一种方式是转导,即由细菌病毒介导的遗传物质交换。噬菌体(bacteriophage)常简称为phage,是一类感染细菌的病毒,其首先附着于宿主细胞表面,随后将其DNA注入细菌细胞内。接着,它会降解宿主细胞自身的DNA,并复制其病毒基因组,同时劫持细胞的合成机制以产生大量病毒蛋白。这些噬菌体蛋白随后会自我组装,并将噬菌体基因组包裹进衣壳,形成多个子代噬菌体。然而,由于DNA包装机制的保真度较低,偶尔噬菌体会将细菌DNA片段包装进噬菌体衣壳中。在诱导宿主细胞裂解后,子代噬菌体被释放;当此类噬菌体感染另一个宿主细胞时,便会将其前一个宿主的DNA片段转移进去。该DNA片段随后可能发生重组并永久整合到新宿主的染色体中,从而实现两个细菌之间的基因转移。
在实验室中进行噬菌体转导需要以下要素:含有目的基因的供体菌株、缺乏该基因的受体菌株、能够感染这两种菌株的噬菌体,以及一种用于筛选转导细菌的方法。在大多数情况下,该方法为一种选择性固体培养基,它能够支持转导细菌的生长,同时抑制未转导细菌的生长。实验开始时,将含有目的基因的供体菌株接种于液体培养基中进行培养。当所有细菌均处于对数生长期并活跃分裂时,向培养物中接种目标噬菌体。经过三至四小时的孵育后,几乎所有细菌均已裂解并释放出噬菌体颗粒,此时将供体噬菌体裂解液接种至新培养的受体菌株培养物中。短暂孵育一小时后,培养物中应包含转导与未转导细菌细胞的混合物,通过将部分菌液涂布于适当的选择性固体培养基上来筛选转导细胞。经过进一步孵育,转导细胞应生长并增殖形成可见菌落。随后可挑选这些菌落,利用多种方法进一步验证转导是否成功,例如菌落PCR、DNA测序或定量PCR。
开始实验操作前,应穿戴适当的个人防护装备,包括实验服和手套。接着,使用70%乙醇对工作区域进行消毒,并擦拭台面。
此后,准备三份每份一毫升的 LB 盐溶液。现在,通过将 100 微升的大肠杆菌加入含有五毫升 LB 培养基并添加 500 微克氨苄青霉素的 15 毫升锥形管中,制备供体菌株培养物。然后,在 37 摄氏度下振荡培养过夜,通气并以 220 转/分钟振荡。第二天,从摇床培养箱中取出培养物前,先用 70% 乙醇擦拭工作台面。接着,通过将 10 微升供体菌液加入 990 微升添加了盐溶液的新鲜 LB 培养基中,将过夜培养物按 1:100 稀释。
将细菌稀释液在37摄氏度、220 rpm振荡和通气条件下培养两小时。当细胞达到对数生长期初期时,从培养箱中取出培养物,向其中加入40微升P1噬菌体,再次孵育。继续监测细胞一至三小时,直至培养物发生裂解。随后,向裂解液中加入50至100微升氯仿,通过涡旋振荡混合。然后,离心裂解液以去除碎片,并将上清液转移至新的无菌管中。向上清液中加入几滴氯仿,于4摄氏度保存,保存时间不得超过一天。
开始转导实验时,取1毫升受体菌株培养物。接着,将100微升供体噬菌体裂解液转移至1.5毫升微量离心管中,在37摄氏度下敞盖孵育30分钟,使残留的氯仿充分挥发。在供体噬菌体裂解液孵育的同时,通过轻柔离心收集受体菌株细胞。弃去上清液,将菌体沉淀重悬于300微升含有100毫摩尔硫酸镁和5毫摩尔氯化钙的新鲜LB培养基中。
接下来,通过在微量离心管中混合100微升受体菌株和100微升供体噬菌体裂解液,建立转导反应体系。然后,通过混合100微升受体菌株和100微升含硫酸镁与氯化钙的LB培养基,设置阴性对照。孵育后,向两个离心管中各加入200微升1摩尔的柠檬酸钠和1毫升LB培养基,并通过轻柔吹打混匀。随后,当离心管孵育1小时后,通过离心轻轻沉淀细胞。
离心后,弃去上清液,并将沉淀的细胞重悬于含有100毫摩尔柠檬酸钠的100微升LB培养基中。涡旋混匀溶液,然后将全部转导样品移至含1倍氨苄青霉素的LB琼脂平板上。最后,将阴性对照细胞混合物的全部体积移至不含氨苄青霉素的LB琼脂平板上。将平板在37摄氏度下过夜培养后,使用无菌移液器吸头从转导平板上挑取三至四个菌落,并划线接种到一个新的含有1倍氨苄青霉素和100微升1摩尔柠檬酸钠的LB琼脂平板上。对阴性对照样品,采用相同的涂板方法,接种至另一含有仅100微升1摩尔柠檬酸钠的LB琼脂平板上。随后,将平板在37摄氏度下过夜培养,以使不含噬菌体的菌落生长。
第二天,用70%乙醇擦拭实验台面,然后从培养箱中取出培养板。使用无菌移液器吸头,从转导平板上挑取三个菌落,并将每个菌落分别加入含有5毫升LB培养基的独立离心管中。接着,从阴性对照平板上挑取三个菌落,加入另一管含有5毫升LB培养基的离心管中。将各培养物在37摄氏度下振荡培养过夜,振荡速度为220 rpm,并保证通气。按照之前演示的方法对实验台面进行消毒后,使用DNA小提试剂盒,根据制造商说明书从每种培养物中提取4.5毫升的DNA。然后,用35微升无核酸酶水洗脱DNA,并使用实验室分光光度计测定其浓度。最后,将两种细菌培养液剩余的0.5毫升分别与0.5毫升100%甘油混合,制备甘油保存菌种。
为确认转导效果,首先配制两份用于24次qPCR反应的qPCR预混液。对于第一份预混液,在微量离心管中加入150 µL qPCR缓冲液混合物,并分别加入12 µL针对氨苄青霉素抗性基因设计的正向引物和反向引物。接着,通过在微量离心管中加入150 µL qPCR预混液,再分别加入12 µL针对看家基因设计的正向引物和反向引物,配制第二份qPCR预混液。
每个qPCR反应中,将来自各反应的100微克实验DNA与14.5微升qPCR预混液混合。接下来,按照之前演示的方法准备其余反应。将反应体系转移至预热至94摄氏度的热循环仪中,然后启动程序。最后,利用qPCR生成的循环定量值(Cq值)计算氨苄青霉素抗性基因的标准化转导效率。
将目的基因在各阴性对照和转导样本中的循环定量值(Cq值)进行列表统计。较低的Cq值,通常低于29个循环,如本示例中的转导样本所示,表明目标序列的含量较高。
此处也列出了一个看家基因,用作上样对照,以标准化每个反应中的DNA量,并作为阳性对照以确保qPCR正常工作。只要上样量相同,该看家基因在每个样本中的检出率相对一致。
接下来,为了计算每个样本的 ΔCq 值,需用每个样本的目标基因 Cq 值减去其看家基因的 Cq 值。例如,第一个阴性对照的 ΔCq 值为 13.54。然后,利用此处所示公式,根据该值计算每个样本的标准化转导效率。最后,可计算出每组样本的平均标准化转导效率。
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Q1: How does a bacteriophage transfer DNA between bacterial cells?
A bacteriophage attaches to a bacterial cell surface and injects its DNA, then replicates its genome while hijacking the host's machinery. Due to low-fidelity DNA packaging, the phage occasionally packages bacterial DNA fragments into its capsid. When this phage infects another cell, it transfers the bacterial DNA, which can recombine and integrate into the new host's chromosome, mediating gene transfer between bacteria.
Q2: What are the key requirements for performing phage transduction in the laboratory?
Phage transduction requires a donor strain containing the gene of interest, a recipient strain lacking it, a phage capable of infecting both strains, and a selective growth media that supports transduced bacteria while inhibiting non-transduced ones. The selective media is essential for screening and identifying which bacterial cells successfully received the transferred genetic material.
Q3: Why is chloroform added to the phage lysate during preparation?
Chloroform is added to the phage lysate to inhibit bacterial growth and preserve the phage particles. After centrifugation removes debris, the supernatant containing phage is treated with chloroform and stored at four degrees Celsius for no more than one day, ensuring the phage remains viable for the transduction procedure.
Q4: What role do magnesium sulfate and calcium chloride play in transduction?
Magnesium sulfate and calcium chloride are added to the recipient strain suspension to enhance phage attachment and DNA transfer efficiency. These divalent cations stabilize the phage-bacterial cell interaction, increasing the likelihood of successful transduction by promoting optimal conditions for DNA uptake by recipient cells.
Q5: How does qPCR confirm successful transduction of the ampicillin resistance gene?
qPCR measures cycle quantification (Cq) values for the ampicillin resistance gene and a housekeeping gene. Low Cq values below 29 cycles indicate high target sequence amounts in transduced samples. Delta Cq values normalize results against the housekeeping gene, and normalized transduction efficiency is calculated to confirm the gene transfer occurred in recipient cells.
Q6: What is the purpose of the negative control in phage transduction experiments?
The negative control combines recipient strain with LB medium containing magnesium sulfate and calcium chloride, but without phage lysate. This control undergoes identical steps as the transduction reaction, allowing researchers to verify that any ampicillin resistance detected in transduced samples resulted from phage-mediated gene transfer, not contamination or spontaneous resistance.
Q7: Why is sodium citrate used during the transduction procedure?
Sodium citrate is added after the one-hour incubation period to chelate divalent cations and halt phage-mediated DNA transfer. This stops the transduction reaction at a defined time point, preventing continued gene transfer and allowing researchers to accurately assess the efficiency of the transduction event that occurred during the incubation window.