October 02, 2013
By: Denise Leo, Global Brownie Ambassador
Category: , , , , ,
This recipe is dangerously simple. Three ingredients. Hardly any effort. No dirty mixing bowls. And you end up stuffing your face with an amazing, tender, crispy, chewy, rich, and memorable dessert. I promise that you’ll want to make this one again and again and again.
INGREDIENTS
2 packages of Nestlé cookie dough (Ultimate Dark Chocolate Delight, in the refrigerated aisle, you know: the “break-n-bake” kind)
Optional: A pinch of flaked salt (like the kind used to line a margarita glass)
5 Caramello candy bars
DIRECTIONS
Coat muffin pan liberally with butter-flavored cooking spray.
Break both packages of cookie dough along the pre-set marks. That will make 24 separate pieces of chocolate cookie dough.
We want big, thick brownie snacks, bigger than just one cookie. So I want you to smoosh together 1 and 1/3 cookies.
Grab that monster-sized bit of dough and put it into a greased muffin tin. (You’ll end up with 18 total.) I liked the effect of having the pretty chocolate chips showing on top. See photo.
Take a pinch of flaked salt and sprinkle a tiny bit on top of each brownie (optional). Bake at 350 F for about 18-20 minutes. They will be a little puffy, but not like a muffin. You can tell with a toothpick if they’re close to done. The toothpick should come out nearly clean. Either of these two toothpicks would suggest your brownie is done.
Remove the brownies from the oven. For each brownie, the puffiness will go down as it starts to cool. Mine were about 1 inch tall. Allow them to cool for a few minutes. (Set a timer so you don’t forget!) After 4-5 minutes of cooling, it’s time add your Caramello squares. If you add the Caramellos too soon, the delicate squares will melt and look saggy. After the short cooling time, break the Caramello bar into squares, and insert a square partway down below the still-soft crust of each brownie.
Yu-u-u-u-u-u-um.
Leave your ooey, gooey, chocolate-y Caramello Brownies in the muffin pan for at least another 15 minutes until they cool and set up a bit. Carefully remove your Caramello Brownies from the pan and then hope they last long enough for your guests to arrive.
June 25, 2013
By: Denise Leo, Global Brownie Ambassador
Category: , , , , , , , ,
This is the FIRST CUPCAKE for my site, and I did NOT make this decision lightly… in truth, once I had the idea for this citrus-y, lemon-y, lime-y dessert, I knew that the Mountain Dew Cupcake simply HAD to be created. Just look at it. It’s like the headline should be “Cupcake Escapes from Kitchen and Rules the X-Games.”
BAM! Yeah, THAT just happened, right in your mouth.
CITRUS BATTER INGREDIENTS
1 box lemon cake mix (I used Duncan Hines Lemon Supreme)
One 6-ounce cup lime yogurt (I used Yoplait Key Lime Pie flavor)
1/3 cup oil
3/4 cup water
MOUNTAIN DEW SWIRL INGREDIENTS
Two 12-ounce cans
(about 3 cups)
Juice of 1 lime (about 3-4 Tbsp)
2 Tbsp cornstarch
3/4 cup of the citrus batter, above
14 drops green food color
3 drops yellow food color
CITRUS FROSTING INGREDIENTS
One tub lemon frosting (I used Duncan Hines Creamy Lemon)
3 drops green food color
2 drops yellow food color
3/4 cup sifted powdered sugar
DIRECTIONS
Put on your shin guards and crash helmet. Keeping one hand on the handlebar of your BMX, use the other hand to simultaneously pour both cans of Mountain Dew into a medium saucepan. You will start with about 3 cups of liquid in the pan.
We’re going to reduce that Mountain Dew down to its intense, dewy essence. There are no rules on how to reduce Mountain Dew, so I made ‘em up: Pow! Boil that bad boy on high heat for about 15-20 minutes until about 3/4 cup of liquid remains.
During the 15 minutes when that’s boiling down, squeeze about 3-4 tablespoons of lime juice into a small bowl and mix in 2 tablespoons of cornstarch. Set that aside for later.
Check your boiling pan. When you have about 3/4 cup of molten Mountain Dew left, it’s time to move forward.
It may be hard to judge how much is in the pan, so you can use a heat-safe Pyrex glass measuring cup to check. That’s what I did.
Keeping the pan off of the heat, return The Dew to the saucepan, except for about 1/8 of a cup, which you will mix right into the cornstarch mixture. The goal is to get the cornstarch mixture warmed up a little.
Add cornstarch mixture to the pan, whisking it quickly, like you’re pedalling up an extreme hill on the BMX course. Don’t worry if you get flecks of lime goo on your face and apron — pretend it’s the mud you’re flinging onto the competitors behind you.
Return the mixture to medium or medium-high heat for just another minute or so, until it thickens up to the consistency of pudding. It will look a lot like lemon meringue pie filling at this point. This, my MD friends, is the basis for your Mountain Dew Goo. Pour it into a medium-sized bowl and set it aside while you make the cake batter.
Put all 5 of the batter ingredients into a large bowl and mix with an electric mixer for 2 minutes on medium speed.
It should be thick, smooth, and creamy. Don’t eat it.
Take about 3/4 cup of the lemon batter and add it to the bowl of the Dew Goo.
Mix together and then add 14 drops of green food coloring.
Getting close. Now add 3 drops of yellow food coloring. Mix until the color is even. You can tweak it to be greener or yellower based on your personal preference. (I found this particular shade to be a nice balance against the yellow of the batter. Just one DewGirl’s opinion.)
Set the green goo aside while you fill all your muffin cups about 1/4 full with the yellow batter. I used foil muffin cups in silver and green, left over from C they were just so Dew-like.
On top of that layer, add an edge-to-edge layer of the green Mountain Dew Goo. This layer should bring each muffin cup up to about half full.Be sure to spread the green from edge to edge so you can’t see any of the yellow batter.
Top off each cupcake with a final layer of the yellow batter, making sure to cover the green entirely. I found the easiest way to do this was to spoon a circle of batter around the outer edge of the cup and then cover over the middle. Each cup should be pretty full, about 3/4 to 5/6 full.
Bake at 350 F for 17-21 minutes, until a toothpick test reveals a nicely raised (but still slightly moist) and spongy-set cake. During baking, something wild happens to that green layer, which gets all loopy, swirly and tossed around as the batter rises and expands. You’ll see what I mean when you take your first bite.
Set aside to cool completely. In the meantime, mix your frosting. Put the pre-made lemon frosting into a medium bowl. Mix in 3 drops of green and 2 drops of yellow food coloring. Feel free to tweak until you get the Mountain-Dew color that YOU like.
Add 3/4 cup of powdered (or confectioners) sugar. Mix until it’s completely lump-free.
Sprinkles make everything better. My opinion: These Mountain Dew Cupcakes just do NOT look extreme without some garish garnish on top. POW! Now THAT’S better!
You don’t HAVE TO refrigerate these, but they are so much more refreshing when you do. Chill off on a summer day with one of these bad boys. YEAH!
Keep your eyes on the prize: the one, the only, Mountain Dew Cupcake!!
May 24, 2013
By: Denise Leo, Global Brownie Ambassador
Category: , , , , , , , ,
You’ll be so dang pleased with yourself when you make this de-freakin’-licious recipe, so just get on it right now: It’s time to introduce your mouth to the fantasy-revelation of bread pudding blondies with a homemade whisky-cream sauce with toasted pecans. (The whisky is optional, but highly recommended.)
I’m salivating as I type this. Seriously.
The amazing thing is that it all starts with a charmingly affordable mini-box of
yellow cake mix! Yep, that’s what I said. It’s probably less than a buck at your local market. Ain’t it cute?
INGREDIENTS – BLONDIES
8-9 cups stale bread, cut into cubes (I bought a 24-ounce sourdough loaf and used 3/4 of it)
1 small box Jiffy yellow cake mix (9 ounces, or about 1 1/2 cups)
3/4 teaspoon ground cinnamon
3/4 teaspoon ground nutmeg
1/3 cup oil
1 Tbsp rum flavor or rum extract
3 1/2 cups milk
3/4 cup fresh, soft raisins
INGREDIENTS – CREAM SAUCE
1 1/2 cups (12 oz.) Half & Half (or whipping cream, if you want it richer)
1/3 cup sugar
2-3 Tbsp corn starch
2-3 ounces whisky (or you can substitute 1 Tbsp rum extract + 1 tsp vanilla)
2 Tbsp butter or margarine
3/4 cup toasted pecans, coarsely chopped
DIRECTIONS
Cut your stale bread into about 1-inch cubes until you have about 8-9 cups. (If your bread isn’t stale, cut into cubes and expose them to air for a few hours. Stale bread does a better job of soaking up your golden delicious batter.)
In a large mixing bowl, blend together the dry cake mix with the ground cinnamon and ground nutmeg. I like to use a fresh nutmeg it makes my kitchen smell so good!
Add the eggs, oil, rum extract, and milk into the mixing bowl. Using an electric mixer, beat for about 2 minutes on medium. This is your blondie batter.
Grease the bottom and sides of a 9×13-inch baking pan. Put all your bread cubes into the pan and then pour all of the blondie batter over top, evenly distributing the batter so the bread will soak it up, leaving no overly dry bread cubes.
While the bread is soaking, sprinkle the top with raisins. It’s looking good, isn’t it?
Bake at 350 F for about 25-30 minutes. To test for doneness, try the toothpick test. Insert the toothpick into the center and look at how wet or dry the toothpick looks. If it’s glossy wet, it needs more time. If some small, moist pieces cling to the toothpick, the blondies are done (see photo below).
The goal is to pull the blondies from the oven before the toothpick comes out completely clean and dry, which would indicate over-baking. (But this recipe has cream sauce, so if you overbake it, you’re still OK; you can pile on the cream sauce and still call it perfectly delicious!) Set the blondies aside and let them cool.
Now it’s time to make the cream sauce. Start by setting aside 1/4 cup of half & half into a small bowl. Pour the remaining 1 1/4 cups of half & half into a medium saucepan.
Add sugar to the saucepan. Mix and cook over medium heat just until it bubbles, stirring occasionally.
While waiting for the sugar and cream to boil, mix 2 T of cornstarch into the 1/4 cup of half & half that you set aside in the bowl. M this is called a slurry.
When the half & half reaches a light boil, pour in the cornstarch mixture and keep cooking on medium heat.
In a minute or two, your cream sauce will be nice and thick. Now add the whisky (or rum extract + vanilla), the butter, and the toasted/chopped pecans. Mix it all together.
If it’s too thick, you can add a few teaspoons of milk, water, or whisky to thin it out.
If it’s too thin for your preference, mix up another slurry of cornstarch-plus-liquid in a one-to-three ratio. You can use 1 Tbsp of cornstarch plus 3 Tbsp of water, or cornstarch plus milk, or cornstarch plus whisky, or cornstarch plus half & half. Add the slurry to the saucepan and cook on medium for another minute or two until it thickens up. (Never add powdered
it can create sticky lumps.)
To serve, cut a piece of bread pudding and heat it in the microwave for 20 seconds. Generously dollop some of that rich, smoky, warm cream sauce right on top. You might even put the cream sauce into a gravy boat and let your guests top their own! So decadent! Store leftover blondies and cream sauce in the refrigerator.
Oh, people. I’m swooning for this guest-worthy recipe. Just look at that photo. How can anyone resist the beauty and the crave-ability of this Bread Pudding Blondie with Whisky-Cream Sauce?
April 10, 2013
By: Denise Leo, Global Brownie Ambassador
Category: , , , , , , , , ,
On May 5, Cinco de Mayo, celebrate with a light, sweet, tart, creamy, lime-bliss concoction that will knock your party socks off: the Mucho Margarita Blondie! PS – It’s one of the easiest recipes on this whole site!
INGREDIENTS
White chocolate macadamia nut cookie dough, 16-ounce package
Whipped topping, 12-ounce tub, thawed
6 ounces frozen-concentrated limeade, thawed (I used half of a 12-ounce can)
1/8 cup to 1/4 cup tequila
Optional food coloring, green and yellow
Optional garnish of flaked margarita salt or toasted flaked coconut
DIRECTIONS
First, thaw both the frozen whipped topping and the frozen-concentrated limeade. While they’re thawing, you can bake the cookie crust and wait for it to cool.
If you love coconut and want to use it as a garnish, shake about 1/2 cup onto a cookie sheet in a thin layer. Bake at 350 F for about 5-7 minutes until it’s golden brown. Watch it carefully, because it will go from perfect to burned in a minute or two. Set coconut aside to use right when serving.
Into a greased 9×13-inch pan, spread the cookie dough, pressing it by hand into a thin layer. (If you want a thick crust, use a 9×9-inch pan and bake longer.)
Bake at 350 F for about 15 minutes, until the edges turn a rich, crispy brown. Set aside until it’s cooled completely.
Now it’s time for the margarita part. This is so fast and easy, you will hardly believe it. In a bowl, add the thawed whipped topping (12 ounces), the thawed limeade (6 ounces), and the tequila (1/8 to 1/4 cup, depending on your taste). Stir with a rubber spatula until combined.
At this point, your margarita topping will be zingy, creamy, and refreshing — but maybe not as pretty as it could be. If you like, try adding 5 drops of green food coloring and 2-3 drops of yellow. You could use a lot more food coloring, if you like, but I tend to use less. Just aim for it to have YOUR idea of a perfect margarita-esque appearance.
Mix your margarita topping until it’s uniform in color. Pour your margarita topping over the cooled cookie crust and spread evenly.
Refrigerate for 30 minutes so it sets up a little. Cut into serving-sized pieces.
If you want to balance the zing of lime and tequila, garnish right before serving with a tiny pinch of flaked margarita salt or with a big handful of toasted flaked coconut.
That’s it! For your Cinco de Mayo party, go with a dessert that’s super simple, definitely delicious, and really refreshing: It’s the Mucho Margarita Blondie!
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淘豆网网友近日为您收集整理了关于Cell-wall Invertases from Rice are Differentially Expressed in Caryopsis during the Grain Filling Stage的文档，希望对您的工作和学习有所帮助。以下是文档介绍：Cell-wall Invertases from Rice are Differentially Expressed in Caryopsis during the Grain Filling Stage Joumal of Integrative Plant Biology 2008,∞(4):466—474Cell-wall Invertases from Rice are DifferentiallyEx! }d。n CaExpressed in a opsis during the Grain Filling Stage(1StateKeyLaboratoryofPlantGenomics,icsandDevelopmentalBiology,theChineseAcademyofSciences,BeijinGraduate School ofthe Cheese Academy ofSciences.Belling 100049,China)Abstractcell-walIInvertase plays an important role (来源：淘豆网[/p-7176437.html])jn sucrosepartitioning between source and ans In higher plants.Tolnvestigats the role of cell-walljnvertases for seed development In dca(Oryza sativa L.)。cDNAs of three putative cell-wal¨nvertase genes OsclNl.OsclN2 and OsclN3 were、Solated.Semi-quantitative reversetranscription-polymerase chainreaction analysis revealed different expression paRems of the three genes in vadous dca ans.In developingcaryopses,they exhibited similar temporal expres(来源：淘豆网[/p-7176437.html])sion panems,expressed highly at the early and middle grain filling stagesand gradually declined to Iow levels afterward.However,the spatial expression pattems of them were very different。withOsClNl primarily expressed in the caryopsls coat。OsC协位In embryo and endosperm,and OsC,们In embryo.Further RNAj仃sjtu hybddization analysis revealed that astrong signal of OsCIN2 mRNA was datected ln the vascular parenchymasurrounding the xylem of the chalazaI vein(来源：淘豆网[/p-7176437.html]) and the aleurone layer。whereas OsClN3 transcript was strongly detected in thevascular parenchyma surrounding椭e phloem of the chalazal vein-cross-cells,the aleurone layer and tha nucellar tissue.These data Indicats that the throe call-walIInvertase genes plementsry/synergeUc roles In assimilate unloadingdudng the grain filling stage.In additiont the cell type-specific expression pattems of OsclN3 In source leaf blades andanthers were also Investigat(来源：淘豆网[/p-7176437.html])ed。and its corresponding physiological roles were discussed.Key words:cell-Oryza set/RNA/seed developmenLWang YQ.Wei XL,Xu HL,Chal CL,Meng K,Zhal HL,Sun AJ.Pang YG,Wu B,Xiao GF,Zhu Z(2008).Cell-wall Invertasss f怕m rice aredifferentially expressed in ceryopsis during the grain filling stage.J.Integr.P『日nf Bi01.50(4)。466074.Available online at
most plant species,assimilated carbon is transported a(来源：淘豆网[/p-7176437.html])ssucrose.Use of sucrose as a source of carbon and energy de-pends on its deavage by either invertase or sucrose synthase,which is crucial for development.growth and carbon partition-Received 6 Feb.2007 Accepted 6 Mar.2007Supported by the State Key Basic Research and Development Ran ofChina().the Innovation Foundation of the Chinese Academyof Sdences(KSCX2.Sw·湖and KSCXl-SW-03),and theProgram forStrategic Scientific AIlianceS between (来源：淘豆网[/p-7176437.html])China and herlands(KNAW-PSA.04.PSA.BD.04 for P.B.F.O..KNAvv_CEP and 04CDP022 forY.X.).’Author for correspondence.Tel:+86(0)10 6487 34∞:Fax:+86(0)10 64髓28∞:E-mail:&zzhu@ic8.∞.cll&.◎2008 In鲥hJ协of Botany.the Chinese Academy of Sciencesdoi:10.”114.08.00641.xing(Sturm and Tang 1 999).Invertase(.#-EC.3.2_1.261is a hydrolase.catalyzing the in'eversible deav-age of sucrose into glucose and fructose(Copenald 1 990).Ba(来源：淘豆网[/p-7176437.html])sed on their subcellular locations.plant invertase can bedassified into cytosolic invertase.vacuolar invertase and cell-walI(extracellular)invertase(Sturm 1999).Cell-walI.nvertaseis aglycosylated forrn with an acid pH optimum and a basicisoelectdc point.which allows it to ionically bind to the cell walIvia positive charges at Iow pH(Kim et a1.2000).Cell.walI invertase is considered to be one of the key en-zymes involved jn establishing sink strength(来源：淘豆网[/p-7176437.html])s in vanous sinktissues(Sturm and Tang 1999).The importance of cell-walIinvertase to regulate seed development js undedined by mutantanalysis in maize(Zea maya L.).In the maize mutant miniature-1.the Iack of expression of an endosperm-specific call-walIinvertase Incw2 in pedicel and endosperm causes interruptionof photoassimilate transport into developing kemel.and as aconsequenca.the seeds are only one竹h of the normal weight呲肿№“№!芏够矗.至啦,心肌窒,一筘小:乏旷抛(来源：淘豆网[/p-7176437.html])溅器抢帅扑甜w孙扣Ⅲ:§呈 万方数据(Mi¨er and Chourey 1992:Cheng et aI.1996).1n fava bean(V耙,a,a6a L.),a celI—wa¨invertase gene V竹CⅥ,,￣yf is observedtO be expressed at the pre.stOrjlge phase Of seed developmentin the thin-wa¨ed parenchyma of seed cOat,a region knownto be the site of DhotOassimiIate unIOading.and a mOdeIof invertase.mediated unlOading prOcess has been proposed(Weber et aI.1995).The essentiaI functjon of ce¨.wa¨invenasefo(来源：淘豆网[/p-7176437.html])r suppIying carbOhydrates t0 ans has alsO beendemOnstrated by tnInsgenic approaches.ApOplastic expressionof a yeast(,Ia,Dmyc8s ce怕Ws,ae L.)invertase in什letuber of potato(Sc),a门帅舳emsum L.)increases tuber size(Sonnewald et aI.1997),and ectopic expression of a plant∞¨-wa¨invertase in roots of Ara6fdops『s(A厢6,dops,s纳af『ana)Ieads tO earIy们Owe r.ng and an increase in whOle pIant biOmass(von Schweinichen and B0ttner 2005).In carrot(Daucus ca,DfaL.),antisense expression studies show廿1at ce¨.wa¨invertasepIays a cruciaI roIe in sucrOsepartitiOning tO deVelOping tap rOOts(.rang et a1. 1999).Suppressed expression of an anther.spec币c∞ll-wa¨invertase N,仃88 in t佑nsgenic tobac。o(~协“a,’a fa6acu,&L.)resuIts in a block du r|ng po¨en deveIoDment(Goe乜et aI.2001).A¨of these resuIts support the essentiaI^Jndion of∞¨-wa¨invertase in carbon Dart.tionina betweensource and sink 0rgans.New progress has been made in studies On the roIe of ce¨.wa¨invertase in the devel0Dment of cereaI seeds besides thework in maize bed above.1n barley(H0rl,e￡,,n yuf口areL.).tempo阳I and spatial∞ordinated exp陀ssion of ceII.wa¨invertases and hexOse transpO—Brs in can—0pses du—ng theea rIy deVelOping stage suggests a rOIe in carbOhydrate 3uppIy(Weschke et aI.2003).In ri∞,four of the nine membe陷0f theri∞∞¨.wa¨jnvertase gene famiIy have been Observed to be ex-pressed in immature seeds(Hirose et aI.2002:Cho et a1.2005:Ishimaru et a1.2005:Ji et aI.2005).RNA加s圩L,hvb ridizationanaIysis 0f OsC¨~7 suggested its critical roIe In carbohyd陀teSupply for develOping caryOpsis du ring the pre—storage phase(Hirose et a1.2002).However,∞¨typ争specific exp怕ssionsof other membe惜in deveIoping∞ryopses have not beenexamined.TherefO旧,mO怕detailed anaIysis 0f exp他ssiOns 0fOther Oe¨一wa¨invertases wi¨heID t0 eIucidate their rOIes in ncecaryOpsis deVeIOpment.In the present study,ce¨type-speci们cexpressions of 0sC,Ⅳ2 and OsC,f、,3 in deVeIOping caryopses。anthers and,or sour∞Ieaf bIades were invesliqated.ResuItScDNA cIOnIng and sequence anaIysisA560.bp f陷gment was amp|ified lhrough怕Ve晤e t阳nscr.ption—poIymerase chain readion(RT乙PCR)by using degene陀teprime噶designed from the conserved fmctosidase m争tif and cysteine∞taI”Ic site 0f known∞¨-wa¨invertases(Stu仃n 1 999).Sequence of this cDNA fragment showed 80%Ce¨一wa¨lnVertases in DeVeIOping CaryOpsis of Rice 467and 83%identities with the cOrrespOnding n喜giOn of the maizeoe¨.1Ⅳa¨invertase genes Jhcw7 and Jhcw2(GenBank accessionnumber【ACC】:AF0501 29 and AFl651 80.旧spedivelv).Af【erthe n∞genome sequence was pub¨shed(Yu et aI.2002)。thispartiaI cDNA sequence was used tO search the rice genOmesequence,and three high hOmOIOqOus sequences were iden—ti仃ed,shOwing 98.5%,72.7%and 65.3%identity tO this partiaIcDNA sequence and IOcated at contiauOus DNA sequences撑1 3373,撑35123 and群7732(http:,,btn..cn,nce).respectiveIy.Fu¨-Iength c亡INAs of the three putative ce¨-wa¨in-vertases were isOlated by F≈T—PCR using gene—speci行c p—mersand FtNA仃om 2-1Ⅳeek-OId seed¨ngs.c亡INAs 0f the three genescOntain ORFs 1 734。1 797 and 1 761 bD in lenath,respectiveIy.They show a high degree Of hOmOIOgy to each Other and t0 OtherhOmOIOgs from maize,barley and wheat at both the nucleotideIeveI and the aminO acid IeveI.with highest hOmology to themaize,仃cw7.,门cw2 and,ncw3(GenBank ACC:AF050129.AFl 651 80 and AF043346),respectively.The three putatjven∞ce¨-wa¨invertase genes were∞rrespOndingIy named as0sC,~7,0sC,~2 and osC,N3(GenBank ACC:AY34231 9。AY340072 and AY342320,respectivelv).Analysis of the deduced prOteins of C)sC,~7.OsC,^f2 andOsC,~3 revealed many typi∞I features of∞¨一wa¨inve广tases。incIuding the conserved卢-fnJc【osidase motif(NDPNG,A),the∞taI”ic cysteine(MWECP,V)site,a basic isoeIednc point,a∞nserVed N-gIy∞sylation site and aputative signaI peptideprobabIy陀quired for e疵阳ce¨uIar lo∞Iization of the Drotein(Figu阳1).The旭are another tl『l帕potentiaI N.gIy(Ⅺsylation sites(NXS门-)Io∞ted at the same positions of the th怕e genes.and two speci仍∞¨y located in OsClN3(Figure 1).Comparingthe cDNA sequen∞s and genomic DNA sequen∞s(NationaICenterfor Biotechnology lnforrnation fNC:B11 database)reveaIedthat the three genes are anized intO six intrOns andseven exOns.and∞ntain a 9-bp exOn 2.encoding three aminOacids(DPN)of the∞nserved脚dDsidase mot讦(NDPNG,A)(Stu丌n 1999).ExpressiOn of 0sC,~f,0sC删2 and 0sC^fV3 in di仟erent—ce ansTissue.specj行c exDressiOns of 0sCJ~,.osC,~2 and OsC,^『3were investigated by semi—quantitative RT—PCR.As shOwn inFigure 2.the three genes show different expressiOn pattemsin va riOus tissues/brgans.OsCf~7 is expressed in a¨sink andsour∞ans tested at very diffb怕ntIeveIs.strongIyexpressed in sink Ieafblades.shoOts and roots ofseed¨ngs。andweakly expressed in source Ieaf bIades。Ieaf sheaths.upper-mOst.ntemodes and anthe阽.0sC,~2 is specm∞IIV exp陀ssedin sink tissues,6rgans.expressed highly in sink Ieaf blades。uppermOst intemodes and anthe倦.and IOwIy in sh00ts androots of seed¨nqs.but is not expreSsed in sOurce Ieaf bIades andIeaf sheaths.0sC,~3 is mOre∞nstitutiveIV expressed in a¨sinkand sOurce“ans tes“翊,with its hiahest exDressiOn in
万方数据468 Journal of Integrative Pfa小Biology V01.50 No,4 2008Figure parison of deduced protein sequences of OsCINl,OsCIN2 and OsCIN3The alignment analysis was carried out with programs Clustal W and BOXSHADE.Conserved pfnJctosidase motif(NDPNG/A),cysteine camlyticsite(MWECP)and N-giycosylation s№are boxed.Another two potential N-gIycosylaUon sites conserved in OsCINl.OsCIN2 and OsCIN3 are doubleundedined,and two sl呲..mc for OsCIN3 are underlined.Arrow indicates the potential signal poptide deavage site.anthers.and relatively high-level expressions in sink Ieaf blades.source Ieaf blades and Iear sheaths.However.in ovaries anddeveloping Caryopses.all of the three genes exhibit broadlysimilar expression paffems.expressed at their highest IeveI.novanes.and stilI at relatively high·levels in caryopses at 4 dafter flowenng(DAF)and 7 DAF.Expressions of OsCfNl andOsC,N3 decrease sharply to abarely detectable leveI at 1 5 and25 DAF.while OsC/N2 expression decreases gradually to a stilIrelatively high IeveI at 1 5 DAF(Figure 2B).Further investigation in different portions of caryopses re-vealed different spatial expression pattems of the three genes(Figure 2C).OsCINf was predominantly expressed in cary-opsis coat.very weakly expressed in embryo。and not ex-pressed in endosperm at a11.OsClN2 was expressed highlyin both embryo and endosperm but very Iowly in Caryop-sis coat.Unlike OsC:/N1 or OsClN2.OsCIN3 was expressedstrongly in embryo。weakly in endosperm.and at abarelydetectable Ievel in caryopsis coat.This iS the first report
万方数据Cell—waJJ Jnvertases Jn Developing Caryopsis of Rice 469Figure 2.Analysis of the expression patterns of OsClNf。OsClN2 and OsClN3 by semi-quantitative reverse transcription·polymerase chain reaction(RT·PCR).(A)Expression in ans.including shoots and roots from 2-week-old seedlings.sink leaf blades and source leaf blades fnom rice plantsUlledng,leaf sheaths of the flag leaves and uppermost intemodes from plants heading.(B)Expression in ans.induding anthers and ovanes 1—3 d before pollination,caryopses at 4.7.1 5 and 25 d after flowenng(DAF).(C)Expression in caryopsis coats.embryos and endosperms of caryopses mixed fnDm 12 lo 15 DAF.Amplification of RAcl cDNA was used as anendogenous contr01.Reproducible results were obtained仟om three independent experiments.Amplified products were separated on 1%agarose gels.Figure 3./n s/tu mRNA Iocalization of OsC,￣3 jn transverse sections of source Ieaf blades and anthereHybddizaUon signal is visible as blue-purple precipitate.(A)Source leaf blade hy晰dized with an osCW3 antisense pmbe.(B)Source lear blade hybridized with an osCIN3 sense probe.(C)Anther hybridized with an OsC,Ⅳ3 antisense probe.(D)Anther hybndized with an OsCIN3 sense probe.ep,mc。po,v。vein.Bars.0.1 mm.of cell-wall invertase expression in developing embryos ofmonocots.Cell type-specific expression of OsClN2 and OsCIN3RNA,n situ hybrydization was carried out on the fuIly openedIeaf blades from 4-week-old seedlings.anthem 1—3 d befo旧poIlination and caryopses at 3.5 and 7 DAF using gene—specificprobes of OsCIN2 and/or OsC,￣3.In the fuIly opened Ieafblades(source Ieaf blades)a very strong signaI of OsC,~3mRNA was detected in mesophyll cells.and weak signalswere detected in epidermal cells and the vascular tissue(Figure 3A).In anthem.a strong signal of OsClN3 transcriptwas detected in the connective vein.and weak signals inpollen grains(Figure 3C).No specific signals were detected、n equivalent sections probed with the OsC|N3 sense probe(F gure 3B,D).A similar result was obtained in anthem forOsCIN2(data not shown).In young Caryopses.difierent cell type·specific expressionpatIems were observed for OsC,~2 and OsCIN3.In caryopsesat 3 DAF.OsC,~2 pt was strongly detected in thevascular parenchyma of the chalazaI vein.weakly detected incross-cells.the nucellar tissue.and endosperm,and almostno signal in pericarp(Figure 4A。C).0sCf~3 mRNA werestrongly detected in the vascular parenchyma surrounding thephloem of the chalazaI vein.cross-cells。nucellar epidermis andendosperm.a Iittle weakly detected in the vascular parenchymasurrounding the xylem of the chalazaI vein,nucellar projectionand nucellus.and weakly detected in pedcarp(Figure 5A.C).Incaryopses at 5 and 7 DAF.strong signals of OsCf~2 transcript
万方数据470 Joumal of Integrative Plant Biology V01.50 No.4 2008Figure 4.in situ mRNA localization of OsClN2 in transverse sections of developing caryopses.Hybridization signal is visible as blue-purple precipitate.(A)Caryopsis at 3 d after flowering(DAF)hybridized with an OsCIN2 antisense probe.(B)Caryopsis at 3 DAF hybridized with an OsClN2 senseprobe.(C)A similar section as(A)at higher magnification.(D)Caryopsis at 5 DAF hybridized with an OsCIN2 antisense probe.(E)Caryopsis at 7 DAF hybridized with an OsCIN2 antisense probe.a,,cross-“.e。i,Iv.1n.Re,np.pe。pedcarp.Bars,0.1 mm.were detected in the vascular parenchyma surrounding thexylem of the chalazal vein and the aleurone layer.weak signalswere detected in the vascular parenchyma surrounding thephloem of the chalazal vein.cross·cells.nucetlar projection andthe inner endosperm,and almost no signals were detected inpericarp and nuceUar epidermis(Figure 4D.E).Strong signalsof OsCIN3 mRNA were detected in the vascular parenchymasurrounding the phloem of the chalazal vein and the aleuronelayer.Weak signals were detected in the vascular parenchymasurrounding the xylem of the chalazal vein,cross-cells,nucellar
万方数据Cell-wall Invertases in Developing Caryopsis of Rice 471Figure 5.In situ mRNA localization of OsClN3 in transverse sections of developing caryopsesHybddizaUon signal is visible as blue-purple precipitate.㈧Caryopsis at 3 d after flowering(DAF)hybridized with an OsCIN3 antisense probe.(B)Caryopsis at 3 DAF hybridized w胁an osC,^f3 sense probe.(C)The same section as(A)at higher magnification.(D)Cawopsis at 5 DAF hybridized、Ⅳi廿1 an OsCIN3 antisense probe.(E)Caryopsis at 7 DAF hybridized州th an OsC,￣3 antisanse probe.a,.cross-CV.e,i.Iv,n,ne,np,pe.pericarp.Bars.0.1 mm.projection。nucellar epidermis and the inner endosperm.anda very weak signal was detected in the pericarp(Figure 5D.E).Furthermore.a weak signaI of OsC,●忆mRNA and a strongsignaI of OsCfN3 mRNA were detected in the IateraI veins ofcaryopses at 3 DAF(Figures 4A,5A),and the signals turnedweaker at 5 and 7 DAF(data not shown).NO specific signalswere detected in equivalent sections probed with OsC/N2 andOsClN3 sense probes。though some background signals weredetected(Figures 4B,5B).DiscussionCell-wall invertase plays animportant role in development,growth and carbon pamUoning in higher plants.In the present
万方数据472 J0uma,0r fnfegraf腑P『anf 8『0,Dgy VOI.50 No.4 2008studV,the three putative ce¨-wa¨invertase genes showeddifI-erent expressiOn pattems in variOus nce 0rgans.indicat-ing distinctiVe physiOlOgi∞I rOIes du r.ng nce deVelOpment(Fiau怕2).A¨of 0sC,~7.OsC,^,2 and OsC』~3 are activeIy in-vOIved in the deveIOpment of sink Ieaf bIades.OvarIes and yOung∞ryopses.OsC,~’may aIso pIay impor【ant roles in shoots andrDots of seedIings,0sC,~2 in the uppermost internOdes andanthers,and 0sC,fV3 in anthers.sOurce leaf bIades and leafsheaths.ExDression of ce¨-wa¨jnvertase in sink ce¨s of ans has p他viousIy been observed(Kingston-Sm.th et aI.1999).However.in the present study。transcrIDt of a∞¨.wa¨invertase 0sCf/V3 was strongIy detected jn mesOphy¨ceIIs.the sour∞ce¨s of sourceleaves(Figure 3A).and∞opera.tive expressiOn 0f a ha ride transporter 0sJWS丁4 inthese oe¨s was also obserVed(Wang et aI.2007),suggestingefficient ha—de suppIy,prObabIy fbr the prDvision ofrespirato哼substrates to suppon energy demands 0f transpOrt.Furthermo怕。0sCf~3 may aIso have a possjble rOIe in sensingchanges in assimiIate abundance and transducjng these intoaltered pattems 0f gene expression.as preViousIy proposed forinvertase in Ieaves(Kjnqston.Smith et a1.1 999).Both 0sC,￣2and 0sCf~3 activeIy participate in anther development at theIate developmentaI stage by predominant expression in theconnective Vein(Figures 2B,3C).indi∞ting the roIe in assim-“ate unIoading.OsC,^『3(0s『￣V们was ea rIier怕po№d to beImponant for microspo旧deveIopment atthe young microspo帕stage and the early bjnucIeate stage(0¨Ver et aI.2005),thedata in the present study prOvide infOrmatiOn for its activeinvOlvement in anther development at the tri-nucIeate stage.an important stage for anther dehiscenoe.0sC,,V2 may DIay asim¨ar impO—ant rOIe in anther deveIOpment.In deVeloping caryopses。the three genes area¨actjveIyexpressed at the earIy and middIe grain fi¨jng stages.andOsC,^f2 may also be important at the Iate grain fi¨ing stage(Figure 2B.C).However.they aredifrerentia¨y expressed,w-lh 0sC『￣7 prjma riIy exp怕ssed in∞ryopsis∞at.OsCf^陀inembryo and endOsperm。and 0sGf,V3 in embryO.M0怕Over.theoe¨type.speci行c expressjon pattems of C)sC,~2 and 0sCf~3a旧di仟e悖nl from each other(Figu陀s 4,5).and from that ofOsC,￣,(Hirose et aI-2002),but they are a¨strongIy ex-pressed in∞¨s/tissues inv0Ived in assimjIate unlOadina and/ortransport.The strOng expression of 0sC,^『2 and 0sCf~3 int11e aIeurone Iayer and embryo suggests that at Ieast partof the assimiIate are transDorted jnto the仃¨aI“ssues hande transponers。which is further supponed bV∞ordinate expressjon of MO haride t佑nsporte隅inthe same region(Wang et aI.2007).These data indi∞te that0sC,~f,0sC,~2 and 0sCf~3,probabIy together w陡h otherceIl-wa¨inVer【ases,pIementary physiOIOgi∞I巾Ies insucrOsepartitioning t0 deveIoping caryopses.and the oe¨一wa¨inVertase,haride t旧nspOrter pathway is impOrtant inassimiIate suppIy fbr caryopsis deveIOpment in—ce.In addition.the strong expression of OsC:f,V3 in the lateraI Veins suggeststhat the IateraI veins may be invOIved in assjmiIate suppIy du r.ngthe Very earIy stage of∞ryopsis development(Figu怕5A)。which were prevjOusIy thought to pIay nO physiOIOgi∞I roIe ingrain fi¨ing(Krishnan and Dayanandan 2003).Wh¨e in cross-∞IIs,an assimiIatOry Iayer propOsed a roIe in photosynthatessupply fbr the endosperm(Ebenezer et aI.1 990),OsCfN3 maypIay a simiIar rOIe as in the mesophy¨ce¨s 0f sOur∞Ieaves.Acting as a centraI moduIator of both concentra“0n andproportion of sucrose and hexoses(including gIu∞se andfructose),which are nol onIv as osmoticum and substratesin carbOn and energy metabO¨sm and in pOIymer biOsynth争sis,bul also as primary messenge懵in signaI t阳nsduction(Ro¨and et aI. 2002). ce¨.wa¨ invertase pIays a pivotaIrOIe in carbon metabO¨sm.assim¨ate Dartjtioning betweensource and a几s,response tO wOunding and infbc-tion。and contrOl Of ce¨differentiation and plant develOpmenI(Sturrn 1999:Sturrn and Tang 1 999:Roitsch et aI.2003).plex仙nctjons of∞II.wa¨invertase are ca州ed out by amuIti.gene famiIy with membe噶showina diverse expressionregulatjon(Kim et a1.2000:Sherson et a1.2003:Cho et aI.2005).The present sludy demonst陀tes that membe隅of this family a怕∞Ope旧tiVeIy expressed and pIementary physi0109i∞IrOIes in pIant deveIOpment.and that the same gene mayplay very different rDIes in sink and sOUr∞“ssues,0rgans.Conside ring pIex expression negulation mechanismsof pIant ce¨一wa¨invertases. including transcr.ptiOnaI.post-transc riptjonaI and biochemicaI controIs(Sturrn 1 999),^jrtherstudies Ons¨encing Or OVerexpressing of singIe genes mightheID t0 eIucidate the djs“nct rOIe Of individuaI ce¨-wa¨invertasein rice develOpment.Mater.aIs and MethodsPIant materiaIsRjce pIantS(o,yza sa打l,a L.cv.Minghui 86)were grown underfieId∞nditions.Each caryopsis was marked On the啊0we r.ngday and subsequentIy sampled fb¨owing matu rity.OnIy∞ry-Opses On the first to third primary阳chis b阳nches frDm the tOpof the panicle we阳sampled.TO obtain tjssue sampIes frOm—ce seed¨ngs.seeds were germinated in water for 2 d.andthen transfen。ed 0nt0 wate卜soaked paDertoweIs for g几D、￣th in agreenhou舱under a 16:8 h Iight:da水叫de at 28。C and 24 oC.陀spectiveIy.FOr RT.PCR。sh00tS and roOts frOm 2-week·oldseedIings.sjnk Ieaf bIades and source leaf bIades frDm plants atlhe t¨Iering stage。leaf sheaths of the伺ag IeaVes and upperrnOstintemodes fn)m pIants at the heading stage,anthers and ovanesat 1—3 d before Do¨ination.and caryOpses at 4,7,15 and 25 DAFwere used.For』h s,山hybridization,the仙¨V opened Ieaf bIades竹Dm 4一week.oId seed¨nqs.anthers 1—3 d before伺owerina andcaryOpses at 3.5 and 7 DAF were used.
万方数据cDNA cloningA partiaI cDNA fragment of OsC,NI and fuII.1ength cDNAsof OsClNl.OsC|N2 and OsC|N3 were 1solated by RT-PCRanalysis.using a pair of degenemta PCR primers ording to the conserved卢-fructosidase motif(NPDNG)andcysteine catalvtic s.1e(MWECPD)of known invertases(Stum'1 9991 and gene-specific primers designed in the predicted 57-and 3-untranslated regions of the putative osc|NI.OsCIN2and OsC,￣3 genes.Prlmer pairs included:for degenerateprimers.5'-GGATCAA(T/C)GA(T,(A/T/C/G)AA(T/C)G(G/C1-37 and 5,一(NG)AA(NG)TC(NGITIC)GG(NG)CA(CIT)TCC—CA.37:for OsCINl,57..37 and 57.GCCGGTATATCGCAGATGG.37:for OsC,Ⅳ2。57-.AAGTGTGAGAGC-37 and 57-GCATTGCATCTATCTCTCTC.37:for OsC,~3.57.GAGGTAAGTAGTGTGT丁AGTGG一37 and 57.GGACGGAGAGAGTAGCTAGC.37.For R.r-PCR.totaI RNAfmm 2-week-old seedlings was reverse-transcribed by using theMoloney murine Ieukemia virus(M-MLⅥreverse transcriptase(Promega.Madison。WI,USA),and resultant first strand cDNAwas used as a template for PCR with primers bed puter analysisNucleotide sequences and amino acid sequences wereprimarily analyzed using the DNAStar software(version4.01)(Madison,WI.USA).Sequence homology searchesjn GenBank were Carried out with the BLAST program(.gov,BLAST,).Splice site predictionwas carried out with programs GENSCAN(http:flgenes.mit.edu/GENSCAN.htmll and FGENESH(ry.phtml).Multiple sequence alignment analysis was Car-ried out with programs ClustaI W(http:ll,clustalw/)and BOXSHADE(http://bioweb.pasteur.fflseqanal/interfaces/boxshade.html).The N·terminal signal sequence wasanalyzed using SignalP and P—sort(/).Semi-quantitative RT-PCRTotaI RNA from rice Caryopses was prepared using RNAcon-cert kit(Tianwei Co..Beijing,China)according to the man—ufacturer's instructions.and total RNA from other tissues ofrice plants was isolated by the method of Chomczynski hi f1 987).A¨RNA samples were treated with DNaseI(Promegal to eliminate DNA contaminaUon.For RT-PCRanalysis,4 Pg of total RNA was reverse-transcribed by usingthe M-MLV reverse transcriptase(Promega)according to themanufacturer's directions.Product of the first-strand cDNAsynthesis reaction was used as a template for amplificationin a PCR reaction with primers previously used for the full-Iength cDNAs amplification of OsCINl.OsC,￣2 and OsCIN3or with primers RAcl.P1 f5,-TGACGGAG.37)and RAcl-P2(57.TCTTGGCTTAG-37)specificfor the rice actin gene尺Acl(GenBank Acc:X1 6280)as anCell-wall Invertases in Developing Caryopsis of Rice 473endogenous contr01.The rice RAcl primers were designedin the region passing an81-bp intron to exclude anyinfiuence by genomic DNA contamination.PCR(25 uL totaIvolume)was Carried out using ExTaq polymerase(TaKaRa.Shiga。Japan).The amount of template cDNA and the numberof PCR cycles were determined by preliminary expenmentsto ensure that amplification occurred in the lJnear range andallowed aOod quantification of the amplified products.For PCR.the amplification program consisted of an initial 95。C for 5 minfoIlowed by 30 cycles(for OsCINl.OsCf~2 and RAc7)or 35cycles(for OsC,Ⅳ3)of94。C,for45s:5昏_60 oC,for45s:72。C.for 2 min.and a finaI extension at 72。C for 5 min.Amplifiedproducts were separated On 1%agarose gels and visualizedby Alphalmager(version 5.5)(San Leandro,CA.USA)forsubsequent analysis.Experiments were repeated three timesto obtain similar results.RNA/n situ hybrydizationRNA/n s#u hybridization was Carried out foIlowing the methodof Hirose et a1.(20021 with slight modification.Plant materialswere fixed in 4%(w/v)paraformaldehyde and 0.25%(v/v)glutaraldehyde in 1 0 mM sodium phosphate buffer(pH 7.0)overnight at 4。C.dehydrated through an ethanoI series anddimethylbenzeI series.and finally embedded in Paraplast Plus(Sigma,St.Louis,MO,USA).Sections(10岬)were mountedonPoly-Prep Slides(Sigma),deparaffinized.and digested withproteinase K(1岖j/mL)for 30 rain at 37。C,followed by post-fixation with 4%(w/v)paraformaldehyde in 1 0 mM sodiumphosphate buffer(pH 7.0)for 1 0 min at room temperature.Subsequently.the sections were treated with acetic anhydride.dehydrated through an ethanoI series。and then hybridizedwith the digoxigenin(DIG)-labeled RNA probes(see below)at 50。C overnight in a hybridization buffer.After hybddization.the sections were treated with RNase A at 37。C for 30 min.foIlowed by incubation with blocking reagent and bovine serumalbumin.The hybridized signal was immunologically detectedusing an anti-DIG alkaline phosphatase conjugate(Roche,Basel,Switzedand)with nitro-blue tetrazolium salt and 5一bromo-4-chloro-3-ind0IyphOsphate toluidinium salt as the substrate.The sections were photographed using the LEICA DMER m.-croscope(Wetzlar.Germany).Since most of the coding region of members of the ricecell-wall invertase gene family shares high similarity witheach other,a 203-bp f阳gment of the 3,untranslated re-gion of osclN2 cDNA.a region very specific fot OsCIN2。was amplified through RT-PCR from 2一week-old seedlingsusing primers 5,.GAAATrGAGAGAG户汀AGATG一37 and ATG-3’.and subsequently cloned intopBlueScdptKS+(Stratagene,La Jolla.CA,USA)as a templatefor/n v/tro transcription.For OsC,￣3.a 1 70-bp fragment of the5’region of its cDNA.including a 24-bp 57 untranslated regionand the first 146_bp coding region,which is very specific for
万方数据474 Journal ofIntegrative Piant Biology V01.50 No.4 2008OsCIN3.was cloned into pBIueScrIptKS+as atemplate forin vitro transcription.The specificity of the probes of OsC|N2and OsC『^『3 was confirmed through BLAST searches of theGenBank database(.gov,).The DIG-Jabeled antisense and sense RNA probes were generated withT7 and T3 polymerases respectively according to the protocolof DIG Northem Starter Kit(Roche).AcknowledgementsWe thank Drs Yihua Zhou and Yonghong Wang(Institute ics and Developmental Biology。the Chinese Academy ofSciences)for assistance in mRNA『几situ hybridization analysis,and Drs Zhukuan Cheng and Fang Shi(Institute of ics andDevelopmental Biology.the Chinese Academy of Sciences)forassistance in microscopic analysis.ReferencesCheng WH,Tallerclo EW.Chouray PS(1996).The miniatureI seedlocus of maize encodes a cell wall invertase required for normaldevelopment of endosperm and maternal calls in the pedicel.PlantCeil 8.971—983.ChoJI,Lee SK。Ko 8,KIm HK,Jun SH,LeeYH et 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Cell-wall Invertases from Rice are Differentially Expressed in Caryopsis during the Grain Filling Stage Joumal of Integrative Plant Biology 2008,∞(4):466—474Cell-wall Invertases from Rice are DifferentiallyEx! }d。n CaExpressed in a opsis during the Grain Filling Stage(1StateKeyLaboratoryofPlantGe...
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