我々は感謝ディスカバリーグラントだけでなく、研究ツールとカナダ自然科学工学研究評議会（NSERC）からインストゥルメントの助成金を介してBHRにサポートを認めます。我々はまた、H.織田と例のライブイメージングシーケンスで使用された遺伝資源を提供するためのブルーミントンショウジョウバエストックセンターを認めます。

準備<ol><li>標準的なブドウジュース寒天プレート4の胚を収集する。それは自動化されたショウジョウバエの卵コレクタ（Flymax科学機器（株））を使用すると便利です。同期を使用すると、胚のコレクションが必要な発育段階で胚の適切な数値を得るために実行しなければならないdechorionationsの数を減らす予定上演。 </li><li>顕微鏡のスライドに両面テープの一部を添付してdechorionationのスライドを準備する。テープから裏紙を取り外します。 </li><li>チャンバーのウェル中に収まるように組織片を切断することにより、ライブイメージング室を準備します。ウェル内の組織を配置し、蒸留水で濡らす。 </li><li> 1:1の比率でハロカーボンオイル（ハロカーボン製品社）粘度のシリーズ700とシリーズ56を混ぜる。混合物は、保存され、無期限に使用することができます。 </li><li>カバースリップ（22 X 40ミリメートル、第2号）の表面にハロカーボンオイルのいくつかの小滴を置きます。ボリュームは重要ではないが、20〜40μlの順で指定する必要があります。 </li></ol>手順<ol><li>実体顕微鏡の助けを借りて、どちらかの宝石商のピンセット（5番）や非常に細かいペイントブラシを使用して、グレープジュース、寒天プレートから胚を回収する。胚は、相互に及び鉗子&quot;のヒントに固執する傾向があり、それらは簡単にまとまって集まっている。 </li><li>実体顕微鏡の助けを借りて、軽くdechorionationのスライド上のテープの表面に鉗子の先端に付着した胚の塊を下げる。 </li><li>再び実体顕微鏡を使用して、静かに内側の卵黄膜（卵黄膜が破裂している場合胚がバーストされる）を破裂させずに外側のワックス状の絨毛膜を開いて打開するために鉗子の面で胚を揺らす事や脳卒中。 </li><li>外側の絨毛膜が破裂した後は、絨毛膜から胚をいじめる。dechorionated胚は、鉗子の表面に付着する傾向があります。テープの表面にdechorionated胚を触れないように注意してください。 </li><li>とすぐに胚がdechorionatedなので、すぐにカバースリップ上にハロカーボンオイルの予め調製したドロップに移す。ハロカーボンの油滴の表面にdechorionated胚を運ぶ鉗子の先端をタッチして - これは石油に鉗子から胚をdislodges。 </li><li>油滴の胚の転送を確認してください。これは、黒い背景上で動作し、斜めの角度で設定された光ファイバー照明の光源を使用していることが肉眼で行うことができます。 </li><li>別の胚をdechorionateを試みる前に、鉗子から任意の油をきれいに。油でコーティングされた鉗子はdechorionationをより困難に、絨毛膜の表面上に小さいものがある。 </li><li> Dechorionated胚は、ハロカーボンのオイルで好調です。鉗子や絵筆を使用し、実体顕微鏡を通して見ながら、油滴の底面に胚を押してお好みのポジションに配置。 </li><li>すぐにライブイメージングチャンバーの優に超えるカバースリップを反転。胚は、所望の方向にカバースリップに対して休んでいることを確認してください。油で彼らの浮力のために、胚はカバースリップの下面に対してフロートします。テープでライブイメージング室にカバースリップを修正。テープがカバースリップの端とライブイメージングチャンバーのウェルの端との間のギャップにまたがる地域にテープで穴をカットすることによりチャンバーを換気する。 </li><li>直立蛍光顕微鏡または蛍光実体顕微鏡は、現在のイメージ胚をするために使用することができます。 </li></ol>

<ol>
	<li>Kiehart, D. P., Galbraith, C. G., Edwards, K. A., Rickoll, W. L., Montague, R. A. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Multiple+forces+contribute+to+cell+sheet+morphogenesis+for+dorsal+closure+in+Drosophila.">Multiple forces contribute to cell sheet morphogenesis for dorsal closure in Drosophila.</a> J. Cell Biol. 149, 471-490 (2000).</li><li>Schock, F., Perrimon, N. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Cellular+processes+associated+with+germ+band+retraction+in.">Cellular processes associated with germ band retraction in.</a> , 248-2429 (2002).</li><li>Reed, B. H., Wilk, R., Schock, F., Lipshitz, H. D. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Integrin-dependent+apposition+of+Drosophila+extraembryonic+membranes+promotes+morphogenesis+and+prevents+anoikis.">Integrin-dependent apposition of Drosophila extraembryonic membranes promotes morphogenesis and prevents anoikis.</a> Curr. Biol. 14, 372-380 (2004).</li><li>Wieschaus, E. p., Nusslein-Volhard, C., Roberts, D. B. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=.">.</a> Drosophila: a practical approach. , 199-227 (1986).</li></ol>

我々は、ハンギングドロッププロトコルを呼び出すのライブイメージング解析、のために、ショウジョウバエの胚を作製する新しい方法を説明します。残念ながら、それは倒立顕微鏡を扱う場合ハンギングドロッププロトコルを使用することはできません。このケースで挟み込む技術が（上記の抽象的で説明したように）使用され、胚の圧縮は、懸念のままする必要があります。 <p class="j...

顕微鏡スライド、カバーグラス（22 × 40ミリメートル、2号）、両面テープ、宝石商の鉗子（デュモンスタイル第5号：標準装備し、ハンギングドロッププロトコルを使用して、ライブイメージングのために胚を作成するために必要な材料は、以下の項目が含まれています）、ハロカーボンオイルシリーズ56とシリーズ700（ハロカーボン製品社）、ユーティリティナイフまたは単一のエッジのかみそりの刃、ティッシュペーパー、はさみ、蒸留水、汎用テープ、20〜40μlを配信するための適切なピペット。ライブイメージングプロトコルはまた、カスタムメイドのライブイメージングチャンバーを必要とします。この値は5 mmは標準的な顕微鏡スライド（75 × 25 mm）の寸法にポリカーボネートプラスチックシートを考えるとスライド（20 X 55 mm）を3mmの深いうつ病を作成するにはローターを使って切断することによって調製される。実体顕微鏡（顕微鏡を解剖）と光ファイバ照明用光源へのアクセスも必要です。

the preparation of drosophila embryos for live-imaging using the hanging drop protocol

緑色蛍光タンパク質（GFP）をベースボックスに追加ライブイメージングは​​、そのような組織の形態形成、細胞間接着、または細胞死などの動的過程の遺伝的調節を研究するための強力な手法です。 GFPを発現するショウジョウバエの胚は、容易にどちらかの立体または共焦点顕微鏡を用いて画像化している。あらゆるライブイメージングプロトコルの目標は、このような脱水や低酸素などの有害な影響を最小限に抑えることです。ライブイメージング解析のためのショウジョウバエの胚を調製するための以前のプロトコルは、ハロカーボンのオイルでdechorionated胚を配置し、ハロカーボンガス透過膜とカバーの間にそれらを挟む関与している<sup&gt; 1月3日</sup&gt;。この方法で取り付け胚を通じて、圧縮の導入は、形態形成のあらゆる生体力学ベースの解析のために望ましくない合併症を表します。我々はハンギングドロッププロトコルを呼び出す私たちの方法、、ライブイメージング中の胚の優秀な生存率の結果と胚が圧縮されている必要はありません。簡単に言えば、ハンギングドロッププロトコルは、順番に、湿度の高い室以上の位置に固定されるカバーガラス、から中断されているハロカーボンオイルの滴で胚の配置を含む。ガス交換を提供し、脱水を防ぐことに加えて、この配置は、タイムラプス買収時の位置から流出することを防止するためにハロカーボンのオイルの胚の浮力を利用しています。このビデオでは、ハンギングドロッププロトコルを使用して、ライブイメージングのためのショウジョウバエの胚を収集して準備する方法について詳細に説明します。このプロトコルは、実体顕微鏡あるいは直立化合物の蛍光顕微鏡を用いたイメージングdechorionated胚に適しています。

Watch this Scientific Journal Video about The Preparation of Drosophila Embryos for Live-Imaging Using the Hanging Drop Protocol at JoVE.com

The Preparation of Drosophila Embryos for Live-Imaging Using the Hanging Drop Protocol

ライブイメージング解析のためのショウジョウバエの胚を作製する、シンプルで安価な、そして効果的な方法が提示される。我々のプロトコルは、湿度およびガス交換を提供し、ショウジョウバエの胚を圧縮しません。このメソッドは、実体顕微鏡や正立複合顕微鏡を用いてショウジョウバエの胚のGFPベースのライブイメージングに適しています。

の調製ショウジョウバエ</emハンギングドロッププロトコルを使用してライブイメージングのための>胚

Green fluorescent protein (GFP)-based timelapse live-imaging is a powerful technique for studying the genetic regulation of dynamic processes such as ...

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Research

JoVE Journal

Biology

The Preparation of Drosophila Embryos for Live-Imaging Using the Hanging Drop Protocol

15.1K ビュー.  University of Waterloo. ライブイメージング解析のためのショウジョウバエの胚を作製する、シンプルで安価な、そして効果的な方法が提示される。我々のプロトコルは、湿度およびガス交換を提供し、ショウジョウバエの胚を圧縮しません。このメソッドは、実体顕微鏡や正立複合顕微鏡を用いてショウジョウバエの胚のGFPベースのライブイメージングに適しています。

ビデオ: The Preparation of Drosophila Embryos for Live-Imaging Using the Hanging Drop Protocol

Preparation of Neuronal Cultures from Midgastrula Stage Drosophila Embryos

This video illustrates the procedure for making primary neuronal cultures from midgastrula stage Drosophila embryos. The methods for collecting embryos and their dechorionation using bleach are demonstrated. Using a glass pipet attached to a mouth suction tube, we illustrate the removal of all cells from single embryos. The method for dispersing cells from each embyro into a small (5 l) drop of medium on an uncoated glass coverslip is demonstrated. A view through the microscope at 1 hour after plating illustrates the preferred cell density. Most of the cells that survive when grown in defined medium are neuroblasts that divide one or more times in culture before extending neuritic processes by 12-24 hours. A view through the microscope illustrates the level of neurite outgrowth and branching expected in a healthy culture at 2 days in vitro. The cultures are grown in a simple bicarbonate based defined medium, in a 5% CO2 incubator at 22-24&deg;C. Neuritic processes continue to elaborate over the first week in culture and when they make contact with neurites from neighboring cells they often form functional synaptic connections. Neurons in these cultures express voltage-gated sodium, calcium, and potassium channels and are electrically excitable. This culture system is useful for studying molecular genetic and environmental factors that regulate neuronal differentiation, excitability, and synapse formation/function.

This video demonstrates the preparation of primary neuronal cultures from midgastrula stage Drosophila embryos. Views of live cultures show cells 1 hour after plating and differentiated neurons after 2 days of growth in a bicarbonate-based defined medium. The neurons are electrically excitable and form synaptic connections.

Midgastrulaステージショウジョウバエの胚からの神経培養の準備

This video illustrates the procedure for making primary neuronal cultures from midgastrula stage Drosophila embryos. The methods for collecting embryos ...

生物学

In vitro Differentiation of Mouse Embryonic Stem (mES) Cells Using the Hanging Drop Method

Stem cells have the remarkable potential to develop into many different cell types. When a stem cell divides, each new cell has the potential to either remain a stem cell or become another type of cell with a more specialized function, This promising of science is leading scientists to investigate the possibility of cell-based therapies to treat disease.

When culture in suspension without antidifferentiation factors, embryonic stem cells spontaneously differentiate and form three-dimensional multicellular aggregates. These cell aggregates are called embryoid bodies(EB). Hanging drop culture is a widely used EB formation induction method. The rounded bottom of hanging drop allows the aggregation of ES cells which can provide mES cells a good environment for forming EBs. The number of ES cells aggregatied in a hanging drop can be controlled by varying the number of cells in the initial cell suspension to be hung as a drop from the lid of Petri dish. Using this method we can reproducibly form homogeneous EBs from a predetermined number of ES cells. 

In vitro Differentiation of Mouse Embryonic Stem mES Cells Using the Hanging Drop Method

This video demonstrates how to conduct in vitro differentiation of mouse embryonic stem cells to embryoid bodies using the hanging drop method.

ハンギングドロップ法を用いたマウス胚性幹（MES）細胞のin vitroにおける分化

Stem cells have the remarkable potential to develop into many different cell types. When a stem cell divides, each new cell has the potential to either ...

Live Imaging of Glial Cell Migration in the Drosophila Eye Imaginal Disc

Glial cells of both vertebrate and invertebrate organisms must migrate to final target regions in order to ensheath and support associated neurons. While recent progress has been made to describe the live migration of glial cells in the developing pupal wing (1), studies of Drosophila glial cell migration have typically involved the examination of fixed tissue. Live microscopic analysis of motile cells offers the ability to examine cellular behavior throughout the migratory process, including determining the rate of and changes in direction of growth. Paired with use of genetic tools, live imaging can be used to determine more precise roles for specific genes in the process of development. Previous work by Silies et al. (2) has described the migration of glia originating from the optic stalk, a structure that connects the developing eye and brain, into the eye imaginal disc in fixed tissue. Here we outline a protocol for examining the live migration of glial cells into the Drosophila eye imaginal disc. We take advantage of a Drosophila line that expresses GFP in developing glia to follow glial cell progression in wild type and in mutant animals.

Here we describe a protocol to examine the migration of glial cells into the developing Drosophila eye using live microscopic analysis paired with GFP tagged glial cells.

ショウジョウバエの眼成虫ディスクに付属してグリア細胞の移行のイメージングを生きる

Glial cells of both vertebrate and invertebrate organisms must migrate to final target regions in order to ensheath and support associated neurons.  While ...

Live Imaging of the Zebrafish Embryonic Brain by Confocal Microscopy

In this video, we demonstrate the method our lab has developed to analyze the cell shape changes and rearrangements required to bend and fold the developing zebrafish brain (Gutzman et al, 2008). Such analysis affords a new understanding of the underlying cell biology required for development of the 3D structure of the vertebrate brain, and significantly increases our ability to study neural tube morphogenesis. The embryonic zebrafish brain is shaped beginning at 18 hours post fertilization (hpf) as the ventricles within the neuroepithelium inflate. By 24 hpf, the initial steps of neural tube morphogenesis are complete. Using the method described here, embryos at the one cell stage are injected with mRNA encoding membrane-targeted green fluorescent protein (memGFP). After injection and incubation, the embryo, now between 18 and 24 hpf, is mounted, inverted, in agarose and imaged by confocal microscopy. Notably, the zebrafish embryo is transparent making it an ideal system for fluorescent imaging. While our analyses have focused on the midbrain-hindbrain boundary and the hindbrain, this method could be extended for analysis of any region in the zebrafish to a depth of 
80-100 &mu;m.

In this video, we demonstrate a method by which to analyze the developing vertebrate brain in live zebrafish embryos at single cell resolution by confocal microscopy. This includes the method by which we inject the single-cell zebrafish embryo and subsequently mount and image the developing brain.

共焦点顕微鏡によるゼブラフィッシュ胚の脳のイメージングを生きる

In this video, we demonstrate the method our lab has developed to analyze the cell shape changes and rearrangements required to bend and fold the ...

Preparation of embryos for Electron Microscopy of the Drosophila embryonic heart tube

The morphogenesis of the Drosophila embryonic heart tube has emerged as a valuable model system for studying cell migration, cell-cell adhesion and cell shape changes during embryonic development. One of the challenges faced in studying this structure is that the lumen of the heart tube, as well as the membrane features that are crucial to heart tube formation, are difficult to visualize in whole mount embryos, due to the small size of the heart tube and intra-lumenal space relative to the embryo. The use of transmission electron microscopy allows for higher magnification of these structures and gives the advantage of examining the embryos in cross section, which easily reveals the size and shape of the lumen. In this video, we detail the process for reliable fixation, embedding, and sectioning of late stage Drosophila embryos in order to visualize the heart tube lumen as well as important cellular structures including cell-cell junctions and the basement membrane.

Preparation of embryos for Electron Microscopy of the Drosophila embryonic heart tube

We describe a process for fixation, embedding, sectioning, and imaging of late stage Drosophila embryos for Trasmission Electron Microscopy of the embryonic heart tube. This technique allows for the visualization of the heart tube lumen as well as the basement membrane, which lines the lumen of the heart.

の電子顕微鏡のための胚の調製ショウジョウバエ胎児心臓の管

The morphogenesis of the Drosophila  embryonic heart tube has emerged as a valuable model system for studying cell migration, cell-cell adhesion and cell ...

Live Imaging Of Drosophila melanogaster Embryonic Hemocyte Migrations

Many studies address cell migration using in vitro methods, whereas the physiologically relevant environment is that of the organism itself. Here we present a protocol for the mounting of Drosophila melanogaster embryos and subsequent live imaging of fluorescently labeled hemocytes, the embryonic macrophages of this organism. Using the Gal4-uas system1 we drive the expression of a variety of genetically encoded, fluorescently tagged markers in hemocytes to follow their developmental dispersal throughout the embryo. Following collection of embryos at the desired stage of development, the outer chorion is removed and the embryos are then mounted in halocarbon oil between a hydrophobic, gas-permeable membrane and a glass coverslip for live imaging. In addition to gross migratory parameters such as speed and directionality, higher resolution imaging coupled with the use of fluorescent reporters of F-actin and microtubules can provide more detailed information concerning the dynamics of these cytoskeletal components.

Live Imaging Of Drosophila melanogaster Embryonic Hemocyte Migrations

Drosophila hemocytes disperse over the entirety of the developing embryo. This protocol demonstrates how to mount and image these migrations using embryos with fluorescently labelled hemocytes.

のライブイメージングキイロショウジョウバエ胚性血球移行

Many studies address cell migration using in vitro methods, whereas the physiologically relevant environment is that of the organism itself. Here we ...

Preparation of Developing and Adult Drosophila Brains and Retinae for Live Imaging

The Drosophila brain and visual system are widely utilized model systems to study neuronal development, function and degeneration. Here we show three preparations of the brain and visual system that cover the range from the developing eye disc-brain complex in the developing pupae to individual eye and brain dissection from adult flies. All protocols are optimized for the live culture of the preparations. However, we also present the conditions for fixed tissue immunohistochemistry where applicable. Finally, we show live imaging conditions for these preparations using conventional and resonant 4D confocal live imaging in a perfusion chamber. Together, these protocols provide a basis for live imaging on different time scales ranging from functional intracellular assays on the scale of minutes to developmental or degenerative processes on the scale of many hours.

Preparation of Developing and Adult Drosophila Brains and Retinae for Live Imaging

This protocol describes three Drosophila preparations: 1) adult brain dissection, 2) adult retina dissection and 3) developing eye disc- brain complexes dissection. Emphasis is laid on special preparation techniques and conditions for live imaging, although all preparations can be used for fixed tissue immunohistochemistry.

開発と大人の準備ショウジョウバエ</emライブイメージングのための>脳と網膜

The Drosophila brain and visual system are widely utilized model systems to study neuronal development, function and degeneration. Here we show three ...

Preparing Individual Drosophila Egg Chambers for Live Imaging

Live cell imaging is an important technique applied to a number of Drosophila tissues used as models to investigate topics such as axis specification, cell differentiation and organogenesis 1. Correct preparation of the experimental samples is a crucial, often neglected, step. The goal of preparation is to ensure physiological relevance and to establish optimal imaging conditions. To maintain tissue viability, it is critical to avoid dehydration, hypoxia, overheating or medium deterioration 2. 
 The Drosophila egg chamber is a well established system for examining questions relating, but not limited, to body patterning, mRNA localization and cytoskeletal organization 3,4. For early- and mid-stage egg chambers, mounting in halocarbon oil is good for survival in that it allows free diffusion of oxygen, prevents dehydration and hypoxia and has superb optical properties for microscopy. Imaging of fluorescent proteins is possible through the introduction of transgenes into the egg chamber or physical injection of labeled RNA, protein or antibodies 5-7. For example, addition of MS2 constructs to the genome of animals enables real time observation of mRNAs in the oocyte 8. These constructs allow for in vivo labeling of mRNA through utilization of the MS2 bacteriophage RNA stem loop interaction with its coat protein 9. 
 Here, we present a protocol for the extraction of ovaries as well as isolating individual ovarioles and egg chambers from the female Drosophila. For a detailed description of Drosophila oogenesis see Allan C. Spradling (1993, reprinted 2009) 10.

Preparing Individual Drosophila Egg Chambers for Live Imaging

The Drosophila egg chamber is an excellent model for studying the mechanisms of mRNA localization. In order to capture the dynamic events that underpin the processes of localization, rapid high resolution imaging of live tissue is required. Here, we present a protocol for dissection and imaging of live samples with minimal disruption.

個々の準備ショウジョウバエ</emライブイメージングのための>エッグチェンバース

Live cell imaging is an important technique applied to a number of Drosophila tissues used as models to investigate topics such as axis specification, ...

Live Imaging of GFP-labeled Proteins in Drosophila Oocytes

The Drosophila oocyte has been established as a versatile system for investigating fundamental questions such as cytoskeletal function, cell organization, and organelle structure and function. The availability of various GFP-tagged proteins means that many cellular processes can be monitored in living cells over the course of minutes or hours, and using this technique, processes such as RNP transport, epithelial morphogenesis, and tissue remodeling have been described in great detail in Drosophila oocytes1,2.
The ability to perform video imaging combined with a rich repertoire of mutants allows an enormous variety of genes and processes to be examined in incredible detail. One such example is the process of ooplasmic streaming, which initiates at mid-oogenesis3,4. This vigorous movement of cytoplasmic vesicles is microtubule and kinesin-dependent5 and provides a useful system for investigating cytoskeleton function at these stages.
Here I present a protocol for time lapse imaging of living oocytes using virtually any confocal microscopy setup.

Live Imaging of GFP-labeled Proteins in Drosophila Oocytes

A protocol for live imaging of GFP-tagged proteins or autofluorescent structures in individual Drosophila oocytes is described.

でGFP標識タンパク質のイメージングを生きる<em&gt;ショウジョウバエ</em&gt;卵母細胞

The  Drosophila oocyte has been established as a versatile system for investigating  fundamental questions such as cytoskeletal function, cell ...

Live Imaging of Apoptotic Cell Clearance during Drosophila Embryogenesis

The proper elimination of unwanted or aberrant cells through apoptosis and subsequent phagocytosis (apoptotic cell clearance) is crucial for normal development in all metazoan organisms. Apoptotic cell clearance is a highly dynamic process intimately associated with cell death; unengulfed apoptotic cells are barely seen in vivo under normal conditions. In order to understand the different steps of apoptotic cell clearance and to compare 'professional' phagocytes - macrophages and dendritic cells to 'non-professional' - tissue-resident neighboring cells, in vivo live imaging of the process is extremely valuable. Here we describe a protocol for studying apoptotic cell clearance in live Drosophila embryos. To follow the dynamics of different steps in phagocytosis we use specific markers for apoptotic cells and phagocytes. In addition, we can monitor two phagocyte systems in parallel: 'professional' macrophages and 'semi-professional' glia in the developing central nervous system (CNS). The method described here employs the Drosophila embryo as an excellent model for real time studies of apoptotic cell clearance.

Live Imaging of Apoptotic Cell Clearance during Drosophila Embryogenesis

Here we describe an effective method for studying dynamics of apoptotic cell clearance in vivo. This method employs live Drosophila embryos as a powerful model for monitoring phagocytosis of apoptotic cells using specific labeling of apoptotic cells and phagocytes.

アポトーシス細胞のクリアランスのイメージング中に生きる<em&gt;ショウジョウバエ</em&gt;胚

The  proper elimination of unwanted or aberrant cells through apoptosis and  subsequent phagocytosis (apoptotic cell clearance) is crucial for normal  ...

の調製ショウジョウバエ胚

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Midgastrulaステージショウジョウバエの胚からの神経培養の準備

ハンギングドロップ法を用いたマウス胚性幹（MES）細胞のin vitroにおける分化

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