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Viable Rat-Mouse Chimeras Where Do We Go from Here

Viable Rat-Mouse Chimeras Where Do We Go from Here
Viable Rat-Mouse Chimeras Where Do We Go from Here

and supported by,the phenotypic PCP features of Frizzled/PCP core group genes on one side and that of the Fat/ Dachsous system on the other.Flies carrying mutations in Frizzled/PCP core proteins exhibit defects in PCP throughout the wing.In contrast,the Fat/Dachsous system mainly affects polarity in the proximal half of the wing, as this area strongly depends on cellular realignment and rotation during the switch to the proximal-distal PCP axis. Together,these observations provide an exciting new framework for under-standing the generation of PCP and its relation to new mechanisms that sculpt

the shape of organs in general.

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Viable Rat-Mouse Chimeras:

Where Do We Go from Here?

Davor Solter1,*

1Institute of Medical Biology,A*STAR,138648Singapore,Republic of Singapore

*Correspondence:davor.solter@https://www.sodocs.net/doc/0710885333.html,.sg

DOI10.1016/j.cell.2010.08.021

In a tour-de-force study,Kobayashi et al.(2010)describe the?rst viable rat-mouse chimeras and demonstrate that rat induced pluripotent stem(iPS)cells can rescue organ de?ciency in mice. Rat iPS cells formed a fully functional pancreas when injected into mouse blastocysts lacking the Pdx1gene required for pancreas formation.

Experimentally produced chimeras between different mouse strains(Tarkow-ski,1961)have been an exceedingly useful tool for developmental biologists, contributing to our understanding of the establishment of cell lineages,cell deter-mination,and the development of the immune system and other organs.In this issue of Cell,Kobayashi et al.(2010) dramatically extend the potential of mam-malian chimeras with their report of viable rat-mouse chimeras that can develop to term and become fully functional adults. In their study,Kobayashi and col-leagues relied on previous knowledge but also added a few new wrinkles.They ?rst derived mouse and rat embryonic stem(ES)cells and induced pluripotent stem(iPS)cells using standard methods for the mouse and capitalizing on the

recent isolation of rat ES and iPS cells

(Buehr et al.,2008;Li et al.,2008).Both

mouse and rat cells were tagged with

different?uorescent dyes,allowing the

authors to follow their distribution in the

developing chimeras.The authors wanted

to prove,at least in principle,that xenoge-

neic organ complementation could be

achieved,that is,that donor cells of one

species could rescue a defect in organ

development in a recipient of a different

species.So,as a?rst step,they set out

to produce viable chimeras between rats

and mice,even though many previous

efforts to make such chimeras had failed.

The only viable intergeneric chimera—

that is,a hybrid between animals from

different genera—reported so far is

the geep between a sheep(Ovis aries)

and a goat(Capra hircus)(Fehilly et al.,

1984).

To test the possibility that viable rat-

mouse chimeras could be formed,

Kobayashi et al.(2010)injected?uores-

cently labeled mouse or rat iPS cells into

rat or mouse blastocysts,respectively,

and returned them to blastocyst-compat-

ible pseudopregnant females(that is,

foster mothers of the same species as

the blastocysts).The authors then exam-

ined the resulting fetuses,newborns,and

adults and found evidence of a substantial

contribution of donor stem cells to tis-

sues and organs of the host(Figure1A).

Despite a big contribution of donor cells,

the size of newborn and adult chimeras

(with one exception)was determined by

676Cell142,September3,2010a2010Elsevier Inc.

the species of the host blastocyst.It is not clear whether it is the embryo itself or the uterine environment that determines the extent of chimera growth.To distinguish between these possibilities,one would have to transfer chimeric embryos into the uterus of pseudopregnant females of the same species as the donor stem cells (not the blastocysts).Previous studies suggest that such experiments would fail because of the need for compatibility between the fetal part of the placenta and the uterus (Rossant et al.,1982).

Besides controlling the size and growth of the chimera,the host blastocyst seems to impose additional morphogenetic reg-ulation.The postimplantation develop-ment of normal rat and mouse embryos is very similar,but there are differences in organ morphogenesis.One of the most noticeable differences is the pres-ence of a gall bladder in mice and its absence in rats.In all adult chimeras produced by combining rat stem cells and mouse blastocysts,the resulting ‘‘mouse-like’’chimeras had a gall bladder despite the signi?cant contribution of rat cells to abdominal organs.Reciprocal chimeras were ‘‘rat-like’’and,again,despite a signi?cant contribution from mouse cells to abdominal organs,the gall bladder was absent.These results suggest that cells of the blastocyst inner cell mass possess a ‘‘morphogenetic’’capacity that controls the behavior of injected stem cells at all developmental stages.

This may explain why Kobayashi et al.were able to successfully inject rat stem cells into mouse blastocysts,whereas insertion of the rat inner cell mass into the mouse blastocyst cavity did not result in viable rat-mouse chimeras (Gardner and Johnson,1973).This notion could be tested further using tetraploid comple-mentation (that is,donor ES or iPS cells are injected into tetraploid blastocysts)to produce rat-mouse chimeras (Nagy et al.,1993).Tetraploid blastocyst cells cannot participate in formation of the embryo proper;thus,the resulting fetus (and adult)is derived entirely from the injected cells,whereas the placenta and extraembryonic membranes are derived from the tetraploid blastocyst.It remains to be seen whether this approach could produce a fetus derived entirely from mouse ES cells after their injection into a rat tetraploid blastocyst that then develops in the uterus of a pseudopreg-nant rat female.

A major goal of the Kobayashi et al.study was to determine whether stem cells from a xenogeneic donor mammal could correct a genetic defect in a recipient mammal of a different species.So,in their next set of experiments,the authors in-jected rat iPS cells into recipient mouse blastocysts that lacked the Pdx1

gene,

Figure 1.Generating Rat-Mouse Chimeras

(A)Induced pluripotent stem (iPS)cells were derived from adult mouse and rat cells and were labeled with different ?uorescent proteins.Rat (blue)and mouse (red)iPS cells were injected into reciprocal blastocysts (mouse into rat and vice versa)to produce intergeneric chimeras.From these blastocysts,several chimeras were born and some survived to adulthood.The contribution of injected donor stem cells was observed throughout the body of the host.The size and morphology of the newborn and adult chimeras was determined by the host blastocyst.

(B)Fluorescently labeled rat iPS cells (blue)were injected into normal mouse blastocysts (left)or blastocysts lacking the Pdx1gene (right),which encodes the transcription factor pancreatic and duodenal homeobox 1that is required for pancreas development.Chimeras derived from normal or Pdx1-de?cient mouse blastocysts showed an extensive contribution of rat cells to all tissues.However,in the Pdx1-de?cient chimeras,the entire pancreas was derived from donor rat cells (inset,blue)and was fully functional,including production of insulin by b islet cells.

Cell 142,September 3,2010a2010Elsevier Inc.677

which encodes a transcription factor (pancreatic and duodenal homeobox1) that is essential for development of the pancreas and formation of insulin-producing b islet cells.Although they observed a substantial contribution of rat cells to different organs and tissues,most importantly,the pancreas of the rat-mouse chimeras was composed exclusively of rat cells(Figure1B).Thus,cells derived from rat iPS cells were able to completely rescue the genetic de?ciency of the host mouse blastocyst.These rat-mouse chimeras developed into adult animals with a normal functional pancreas,demon-strating that xenogeneic organ comple-mentation is achievable.This is a remark-able accomplishment.

So where do we go from here?Although human ES and iPS cells offer hope for tissue and cell replacement therapies in the not too distant future,the replacement of complex organs—lung,kidney,liver, gut,and,of course,pancreas—is likely to be much more dif?cult.Several strategies for organ replacement are being tested. These include the growth of organs in vitro with mixtures of different stem cells and biocompatible scaffolds or the gener-ation of‘‘humanized’’pigs engineered to lack certain antigens so that their organs can be used for transplantation in human patients with a reduced chance of immune rejection.Could production of human organs in,for example,human-pig chimeras be an alternative approach? Although production of viable rat-mouse chimeras could be viewed as a?rst step in this direction,as Kobayashi et al. propose,there are huge biological and technical challenges.For example,the

mouse and rat are developmentally very

similar(apart from size),but it is not clear

that chimeras between animals belonging

to different phylogenetic families or orders

would be viable.Indeed,the only attempts

to make such chimeras(between a mouse

and a bank vole)have failed(Mystkowska,

1975).In this experiment,the mouse-bank

vole chimeras were made by aggregation

of embryos;it is possible that injection of

bank vole stem cells into mouse blasto-

cysts,followed by their development

in the uterus of mouse foster mothers

might yield positive results.Successful

chimerism between members of different

orders(the pig and human,for example)

seems very unlikely,and attempts to

produce early postimplantation human-

mouse chimeras have not been encour-

aging(James et al.,2006).Even if we

succeed in developing organ-de?cient

pigs by genetic manipulation and pro-

ducing pig-human chimeras in which the

parenchymal cells of the speci?c organ

are entirely derived from human cells,

immune rejection will still be a problem

because the human organ carried by the

pig will contain pig-derived stromal cells

and blood vessels.

Finally,there are huge legal and ethi-

cal barriers to creating human-animal

chimeras and,indeed,their production is

forbidden in most countries.However,it

is possible that injecting human ES or

iPS cells into a mouse blastocyst and al-

lowing limited(early postimplantation)

development of human-mouse chimeras

would be approved for the speci?c

purpose of testing the differentiation

potential of human stem cells.Yet such

experiments will be complicated,time

consuming,dif?cult to interpret,and,I

suspect,will never become part of the

standard protocols regulating the medical

use of human stem cells.Although xeno-

geneic organ complementation is unlikely

to be a viable strategy for regenerative

medicine,the elegant work of Kobayashi

et al.is a boon for researchers seeking

to better understand the biology of stem

cells and mammalian development.

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Silva,J.,McLay,R.,Hall,J.,Ying,Q.-L.,and Smith,

A.(2008).Cell135,1287–1298.

Fehilly,C.B.,Willadsen,S.M.,and Tucker,E.M.

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Gardner,R.L.,and Johnson,M.H.(1973).Nat.New

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A.H.(2006).Dev.Biol.295,90–102.

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(1)Review feelingsmastered in level 1(happy,sad,hot,cold,tired)。Write the words on the blackboard. Then stand in front of each word and act it out. Ask for volunteers to act out each word. (2)Demonstrate“Can you find my ___?”by giving a student sth of yours and asking him on her to hide it .Close your eyes. Pretend you to have lost the object and lead a dialogue such as:

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Unit 3 under the sea 1.witness (1) v 当场见到,目击 . 在我昨天回家的路上路上我亲眼目睹一场事故。 (2) v表明,说明Her flushed(通红)face she felt. 她通红的脸表明他、她感到非常高兴。 (3) v 为…作证常与to 连用witness to 为某事作证说 He enter the room. 他作证说他看到那个人进入房间。 (4) n (C) “目击者,证人”常与to连用be a witness to 是…的目击者 The police found the witness to the murder case. 警察找到了那个谋杀案的目击者 (5) n (C,U) 证词,证据,证明give witness on behalf of 替…作证 The old man an accused person. 那位老人为被告作证。 2. sort (1) vt 把…分类,拣选 She is . 她正忙于给信件分类。 (2) sort out 整理(好) I 我已把信分好 (3) n 种类,类别 A hammer is .斧头是一种工具。 He’s the sort of person I really dislike. (4)词组sort of 颇为,有几分I feel . 我感觉有点生病了。 all sorts of 相当于all kinds of 3. accommodation (1) 房间,住所The accommodation of this hotel is scare. (不足) (2) 膳宿(常用复数) 我们今晚能找到旅馆住宿吗?Can we ____find accommodations at a hotel__ for tonight? (3) accommodate 向…提供住处,接纳(v) 相关短语:(1) make accommodations for 为…提供膳宿(2) book accommodation at a hotel 在旅馆预订房间 (4) accommodate …to = adapt… to 使…适应 You will have to accommodate yourself to the situation. 4. yell(1)v.叫喊。 (1)yell(out) at sb.朝某人大喊大叫 .她对淘气的孩子大喊大叫。 They yelled at him to stop. (2)n. /叫声,喊声 E.g. a yell of delight / warning 5. throw (1) 投,掷,扔,抛 Throw the ball to your younger brother. He was thrown into prison.被关进… (2) 匆忙或随便穿/脱 He threw a blanket over the injured man. 他匆忙给伤者披上毯子。

人教版高中英语选修七Unit3Underthesea知识讲解Unit3Underthesea语言点

Unit 3 Under the sea 语言点 编稿:牛新阁审稿:王春霞 学习目标 重点词汇 annual witness accommodation opposite yell pause drag depth urge abandon target reflect scare awesome 重点短语 in the meantime be aware of ahead of ... to death help (...)out upside down 重点句型 It takes sb. some time/ ... to do sth.的变化 a time when... 知识讲解 重点词汇 annual 【原句回放】It was a time when the killer whales, or “killers” as they were then called, helped the whalers catch the baleen whales that were on their annual migration. 那个时期,虎鲸(当时被称为“杀手”)帮助捕鲸人在每年须鲸迁徙时捕捉须鲸。 【点拨】annual adj. 1. 每年的,一年一次的 the annual output 年产量an annual report 年度报告 Steel output reached an annual figure of one million tons. 钢的年产量达到100万吨。 2. 按年度计算的 the annual income 年收入the annual rainfall 【拓展】annually adv. 每年地,年度地 witness 【原句回放】I thought, at the time, that this was just a story but then I witnessed it with my own eyes many times. 当时我以为这只是一个故事罢了,但是后来我亲眼见过多次。 【点拨】witness vt.& vi.亲眼看见,目睹,作证; n.见证人;目击者(与of连用) We were witnessing the most important scientific development of the century. 我们正目睹着本世纪最重要的科学发展。 We witnessed great changes in the city. 我们见证了这个城市的巨大变化。 The police found the witness of the murder case. 警察找到了那起谋杀案的证人。 There was no witness at the scene of the accident. 在事故现场没有目击证人。 【拓展】常见搭配:witness to (doing...)为……作证 bear / give witness to 为......作证 in witness of 作为......的证据 a living witness to 活生生的证人 witness stand 证人席(美)=witness box (英) He witnessed to having seen the man enter the building.

小学英语单词(自然拼读整理)

自然拼读词汇 一.a e i o u 的短音 Aa: a n a m a t f a t h a s a nd b a g c a p c a t h a t f a t s a d h a d r a n p a n c a n m a n b a ck th a t th a nk h a ppy st a nd bl a ck p a nda f a mily dr a gon a pple h a ve M a ths a nimal gr a ndma gr a ndpa bl a ckboard ban a na a pple p a rrot a nimal c a rry r a ng st a nd m a tter c a tch Ee: leg g e t b e d y e s r e d t e n h e lp h e llo d e sk s e ven el e ven tw e lve p e n p e ncil e lephant t e nnis pr e sent dr e ss w e st Ii: i s i t i n h i s s i x b i t c i ty th i s b i g sh i p th i n sk i p f i sh m i lk s i n g g i ve w i ndy an i mal sw i m d i nner w i nter an i mal b i scuit c h i ldren

chopst i cks d i ff i cult dr i nk dumpl i ng f i fteen f i fty h i ll k i d l i sten l i ttle l i ve p i cture P i nk s i xteen s i xty sw i m v i sit w i n w i nner s i t g i ve w i nd w i th sk i p m i ss ch i ck f i n i sh f i nger fam i ly s i ster pol i cemen Oo: o x o n o f n o t d o g o n h o t g o t b o x d o ll d o ctor b o dy s o ng p o p pr o ject st o p o range cl o ck fr o g fr o m l o st s o rry b o x h o t l o st dr o p cr o ss b o ttle c o ffee sh o pping Uu: u p b u s m u m b u mp l u nch s u mmer s u nny u nder r u n r u nning l u ck d u mpling h u ndred h u ngry j u mp j u mping m u ch m u m j u st h u rry

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