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Gatekeeping in an inpatient rehabilitation facility to reduce morbidity and mortality due to cardiac disease: screening program using of BNP and ECG Auto-diagnosis.
- Authors
- Type
- Published Article
- Journal
- Nagoya journal of medical science
- Publication Date
- May 01, 2019
- Volume
- 81
- Issue
- 2
- Pages
- 303–312
- Identifiers
- DOI: 10.18999/nagjms.81.2.303
- Source
- Medline
- Keywords
- Language
- English
- License
- Unknown
The Kaifukuki-Rehabilitation Ward (KRW) is a type of inpatient rehabilitation facility in Japan. In the KRW of our institute, mortality and frequency of emergency referrals in 2013 were rather high, 2.6% and 4.3%, respectively. We aimed to investigate the usefulness of an original gatekeeping system to reduce mortality and morbidity from cardiac complications, and to improve the quality of medical care in the KRW. A total of 370 consecutive patients admitted to the KRW of Kobayashi Memorial Hospital between 1 May 2015 and 31 March 2016 were enrolled in this prospective observational study. All patients underwent a screening evaluation in which we defined patients as being screen positive (SC-positive) if they had at least one of 20 diagnostic ECG codes and/or BNP level over 140 pg/dL at admission. A cardiologist provided weekly interventions to those among SC-positive patients who needed cardiac disease treatment during hospitalization. In all, 129 patients were classified as SC-positive (mean age 80 years, 124 [32%] male), and weekly intervention was needed in 28 patients, including start of cardiac medication in 17 cases. Mortality and frequency of emergency transfer due to cardiac disease during hospital stay were 0.3% and 0.3%, respectively. Our gatekeeping system involving a screening evaluation at admission and weekly intervention in selected patients by a cardiologist may be useful in reducing mortality and rate of transfer due to cardiac disease and may improve quality of medical care in KRWs.


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Certainly. So happens.
Prompt, whom I can ask?
I am sorry, it does not approach me. Perhaps there are still variants?
Yes, it is solved.
I think, that you are not right. I can defend the position.


text:
# of partials/manuscripts requested: 2
genre of partials/manuscripts requested: fantasy (1), YA (1)
Unless you were hiding under a rock these last couple days in the publishing world, you will have heard of Harlequin's new venture Horizons. There are plenty of comment threads about it: Absolute Write, Dear Author, etc.
RWA responded by revoking Harlequin's approved publisher status (text on Kristin Nelson's blog). Then MWA got into the mix (text of their statement on Lee Goldberg's site). Harlequin responded by.... changing the name of the venture, which, imo, doesn't address the points that RWA and MWA were raising. Or, as SFWA put it in their statement: "does not believe that changing the name of the imprint, or in some other way attempting to disguise the relationship to Harlequin, changes the intention."
Here's how some other agents feel about it:
* Ashley Grayson - Harlequin Horizons, a mug's game
* Janet Reid - here and here.
* Rachelle Gardner - a self-publishing rant.
Jackie Kessler's version of the author/Harlequin conversation over the last couple days is a must-read!
Some quick definitions:
Self-publishing -- Writer as publisher. The writer undertakes to arrange editing, printing, distribution, etc. without a third party holding any rights or share of the proceeds.
Vanity/subsidy publisher -- A company that publishes books at the author's expense. A vanity press derives its profits from authors. Sometimes provides additional services -- for a fee -- for design, publicity, etc. These fees can generate many thousands of dollars for the press. It's pay-to-play.
Traditional publisher -- Pays an advance/royalty share to authors on the sale of their books. Money flows towards the author. Profits are based on sales. The publishing company's overhead covers editorial, production, distribution, etc.
What these things mean in a query....
If the query mentions a book that has been published, but does NOT mention the publisher, the tendency is to assume that it's either self-published or from a vanity publisher. On some occasions, a google search might turn up the information, but that depends on whether one has the time or inclination to look.
As far as I'm concerned, a book that has been self-published rarely has much impact in a query. I'm not against self-publishing. There are times when it makes sense for the author (see this interview with Wil Wheaton, for example). And, indeed, there are books like The Shack or Eragon, which show it can sometimes lead to more opportunities. And this isn't a new thing either: The Joy of Cooking was originally self-published in 1931 (3,000 copies by a company that printed fancy labels but had never printed a book before). Those success stories are still a decided minority. Then, again, it depends on the author's definition of success and what the author is looking for in the experience. But the same holds true for an agent looking for new clients.
A book that is published by a vanity or subsidy press.... this can be a bit more complex. Many say it's a negative mark on the writer's reputation to have been involved in this approach. Why moreso than self-publishing? Perhaps it's just the sour taste it leaves behind as these types of publishers tend to prey on an author's hopes and use them to their own advantage. In this scenario, it feels like the publishers are making the writers pay for their dreams, often with little hope of any return due to lack of distribution and poor design quality. On top of that, many of them will often take a cut of the profits (sometimes most of it), so the author is not only footing the bill but then paying the publisher a share of the proceeds too. In this case, what motivation does the publisher have to help the author succeed? As for an agent's feelings on this -- well, see those links above, but in most cases it provides a sheen of unprofessionalism, shows a lack of understanding the workings of publishing, and would put the author in the position of starting with a poor hand on a publishing field that is already anything but level.
Would I tell an author not to self-publish? Not necessarily. But I would tell them to employ due diligence and research what it entails and what they are likely to see in return for all their hard work. The author really needs to understand publishing and reaching their readers in order to decide if this is the right path for them to take.


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Dreamspinner Presents
Ethan Hall plans to fill Saturday, December 23, with junk food and bad TV, a day just for him amid the holiday chaos… until his baby sister calls and begs him to go collect a present for her. At the biggest shopping center in Australia. On the busiest shopping day of the year. Hell no. Right?
Ethan's soft heart gets the best of him. He battles through the parking lot, and in the main shopping concourse, he's trampled, elbowed, and bombarded with terrible holiday music. Then he enters hell itself, a specialty store aimed at women… where he meets Ty. They bond in a sea of estrogen and manic shoppers, fighting together to attain freedom, only to find they're not quite ready go their separate ways.
This was a quick little story about finding the silver lining… in a shopping mall! It was cute, witty and sweet. A nice holiday quickie!
4 of 5 stars
Copy Generously Provided for Honest Review


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Rep. Bill PascrellWilliam (Bill) James PascrellHispanic Caucus campaign arm unveils non-Hispanic endorsements Biden rolls out over a dozen congressional endorsements after latest primary wins Juan Williams: Will the GOP ever curb Trump? MORE (D-N.J.) condemned President TrumpDonald John TrumpHealth insurers Cigna, Humana waive out-of-pocket costs for coronavirus treatment Puerto Rico needs more federal help to combat COVID-19 Fauci says April 30 extension is 'a wise and prudent decision' MORE's warning of possible violence if the Democrats take back the House in the fall, calling it "fascism in the flesh."
"This is fascism in the flesh. This is the language of a fanatic that could be lifted from a tyrant's playbook," Pascrell tweeted Wednesday. "This is not a warning but an incitement. Unless we stand united against it."
This is fascism in the flesh. This is the language of a fanatic that could be lifted from a tyrant's playbook. This is not a warning but an incitement. Unless we stand united against it. https://t.co/BUHOBL9auz— Bill Pascrell, Jr. (@BillPascrell) August 28, 2018
The president told evangelical leaders Monday that Democrats will "violently" overturn Republican gains if they win back the lower chamber.
"They will overturn everything that we've done and they'll do it quickly and violently, and violently," the president said.
"There's violence. When you look at antifa and you look at some of these groups — these are violent people," he continued.
Some factions of antifa, a loosely organized group of self-described anti-fascists, have condoned violence as an appropriate response to what they consider fascism. Most recently, some antifa members were arrested Saturday when they clashed with police at a "Blue Lives Matter" rally.
Trump and other Republicans have tried to link antifa with liberals as they try to stave off a "blue wave" in November. A RealClearPolitics average of polls currently shows Democrats with a nearly 7-point lead over the GOP on the generic ballot.
CNN's Don Lemon on Tuesday appeared to defend antifa after the president's comments about the group to evangelical leaders.
"Listen, no organization is perfect. There was some violence. No one condones violence, but there were different reasons for antifa and for these neo-Nazis to be there," Lemon said. "One, racists, fascists, the other group, fighting racist fascists. There is a distinction there."
CNN chief legal analyst Jeffrey Toobin called Trump's criticism of antifa an "appeal to racism," saying the group "is widely perceived as an African-American association."
"Let's be clear also about what's going on here, the theme here is 'I'm Donald Trump and I'll protect you from the scary black people,'" Toobin said.


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Award Winning Sausage! Our Traditional Pork Sausage was named 2018 Gold Coast Sausage King's Best Pork Sausage. Taylor's Fine Foods Manager and Sausage King Paul has perfected the recipe over the years using fresh pork trim that we bone in-store everyday.
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FE teachers cite workload and lack of time for continuing professional development (CPD) as the main barriers to career progression, according to a survey by the National Foundation for Educational Research. Respondents also mentioned a lack of management encouragement, promotion or development opportunities and focused subject-specific CPD. Only 10% said they had found no barriers.
One in three said more on-the job experience, training courses and support of a mentor/coach or line manager were key elements needed to fulfil their career aspirations, thereby fuelling the most common reason given by respondents for working in FE: "Enjoyment in working with young and adult learners, inspiring the next generation and helping them realise their potential."
Just under a fifth of the £5.5m in government grants awarded to colleges preparing for merger has gone to institutions that have failed to merge or have significantly changed their original proposals, reports FE Week. The details appear in a list of some 83 grants made by the end of May this year, published by the Education and Skills Funding Agency. The largest grant for £200,000 supported the creation of a single college in Cheshire from four colleges, although subsequently only two colleges – South Cheshire and West Cheshire – have merged.
Greater student movement in the form of far more transfer agreements between FE colleges running vocational training courses and universities offering first-degree programmes could increase opportunities to access higher education in England, reports TES. Research from UCL Institute of Education/Birkbeck College and US partner Columbia University, states that despite FE colleges catering for some 8.5% of HE students in England, they are rarely mentioned in government reform documents. Currently, vocationally qualified students are more likely to drop out from or simply never attend university. The researchers say much could be learned from the US approach to creating more diverse student bodies, particularly in selective institutions, which understand the many advantages to be gained by students from different social backgrounds living and learning together.
One in three HE providers gaining a 100% student satisfaction rating in this year's National Student Survey are FE institutions, reports TES. Eight colleges (Aylesbury, Boston, Newbury, Seevic, Southampton City, Stanmore, West Lothian and Pembrokeshire) and two sixth-form colleges (Cardinal Newman and Cirencester) are among 30 institutions to earn full-hearted student approval in the annual survey produced by the Higher Education Funding Council for England. More than 300,000 students responded from 530 universities and colleges. A dozen universities, including Oxford and Cambridge, do not appear in the survey because less than half their final-year first-degree students completed the survey questionnaire.


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Sauce Magazine seeks interns for spring 2020
Attention journalism, communications and English students: Sauce Magazine is seeking Editorial Interns for spring 2020.
We need students with a passion for the St. Louis food scene who want to translate that love to print and online media. As a Sauce Editorial Intern, you will:
• Assist Sauce editorial team with the production of the monthly print publication and daily online products. Duties include, but are not limited to, reporting, conducting interviews, writing and contributing to online and print articles, fact checking, assisting with research for upcoming articles, interview transcription, etc.
• Attend occasional events and tastings with the Sauce editorial team, gaining real-world experience as a food journalist.
• Hone your reporting, writing and editing skills with the goal of producing published clips for use in future portfolios
• Assist Sauce Events Coordinator at Food Truck Friday events and Taco Fest.
• Perform other duties as assigned
The Sauce Editorial Intern must have:
• A passion for the St. Louis food scene and the written word
• A working knowledge of AP Style, grammar rules, Microsoft Office and Mac computer systems
• A working knowledge of various social media platforms (Facebook, Twitter, Instagram, etc.)
• Experience conducting phone interviews and writing news articles for print/online publication
• A personable and professional attitude in online, phone and written communication
• The ability to manage his or her time efficiently; should be a self-starter
• A reliable mode of transportation
This internship is unpaid; internship begins in early January and ends in early May. Scheduling is flexible, but the intern must be available 10 to 12 hours a week.
Interested applicants may submit a cover letter, resume, spring semester class and/or work schedule and three to five writing clips to Catherine Klene, managing editor, digital, at cklene[at]saucemagazine.com. All resumes must be submitted no later than Nov. 27. No calls, please.
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Take the Kroger Pick Up Feedback Survey Anyone here like the idea $5,000 to spend at Kroger? If you're currently salivating over the sheer amount of groceries that could buy, KEEP READING. You can now let Kroger know how a recent pick up experience went through the online Kroger Pick Up Experience customer survey. But why I want to take the time out of my busy day to … [Read more...]
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Specific and heritable genetic interference by double-stranded RNA in Arabidopsis thaliana
Contributed by Elliot M. Meyerowitz
We investigated the potential of double-stranded RNA interference (RNAi) with gene activity in Arabidopsis thaliana. To construct transformation vectors that produce RNAs capable of duplex formation, gene-specific sequences in the sense and antisense orientations were linked and placed under the control of a strong viral promoter. When introduced into the genome of A. thaliana by Agrobacterium-mediated transformation, double-stranded RNA-expressing constructs corresponding to four genes, AGAMOUS (AG), CLAVATA3, APETALA1, and PERIANTHIA, caused specific and heritable genetic interference. The severity of phenotypes varied between transgenic lines. In situ hybridization revealed a correlation between a declining AG mRNA accumulation and increasingly severe phenotypes in AG (RNAi) mutants, suggesting that endogenous mRNA is the target of double-stranded RNA-mediated genetic interference. The ability to generate stably heritable RNAi and the resultant specific phenotypes allows us to selectively reduce gene function in A. thaliana.
In Arabidopsis thaliana, reverse genetic techniques for isolating mutants corresponding to known sequences, such as antisense suppression (1–7), cosuppression by overexpression of the target gene (3, 8, 9), targeted gene disruption (10), or the PCR approach of screening for T-DNA insertion libraries (11, 12) have been developed, but are often insufficient and have many unanticipated difficulties. The widespread identification of differentially expressed genes, homologous genes, and interacting proteins have created a need for potent and efficient methods for obtaining their loss-of-function or reduction-of-function mutants.
Double-stranded RNA (dsRNA)-mediated interference with expression of specific genes has been observed in a number of organisms including Caenorhabditis elegans (13–17), plants (18, 19), Drosophila (20, 21), Trypanosoma brucei (22), and a planarian (23). Although the mechanism of RNA interference (RNAi) is not well understood, it seems to provide an effective way to discover gene function in many organisms (24–26).
To investigate the potential of dsRNA interference with gene activity in A. thaliana, we introduced dsRNA-expressing constructs of selected genes with previously defined functions into plants. Gene constructs delivered into plants with Agrobacterium-mediated transformation are stably integrated into the genome of host cells; thus, RNA expression from these constructs in transgenic plants can be persistent and heritable.
In this study, one gene from each of four major categories of genes involved in flower development was chosen, to determine the ability of RNAi to allow functional assessment of genes with diverse developmental functions in flowers. They are the floral organ identity gene AGAMOUS (AG), the floral meristem-size gene CLAVATA3 (CLV3), the floral meristem identity gene APETALA1 (AP1), and the floral organ number gene PERIANTHIA (PAN) (27–30). The phenotypes produced by dsRNAs corresponding to these genes are similar to those of their previously identified reduction-of-function or loss-of-function mutants (31–36). The progeny from fertile RNAi mutants, such as CLV3 (RNAi) and AP1 (RNAi) plants, also showed phenotypes. In addition to high specificity and heritability, a phenotypic series (weak, intermediate, and strong) was obtained from dsRNA interference. Furthermore, in situ hybridization indicates that endogenous target mRNA is decreased in RNAi mutants. Most constructs that are designed to produce only antisense or only sense RNA do not induce interference. Thus, specific and inheritable dsRNA interference may offer a useful alternative to classical reverse genetic screening of mutants in A. thaliana.
Materials and Methods
A summary of DNA constructs is shown in Fig. 1. In p35S∷A-GUS-S and p35S∷A, constructs were ligated to the BamHI and XbaI sites of pCGN1547 (37) into which an 842-bp fragment of the cauliflower mosaic virus 35S promoter and a 253-bp fragment of the 3′ end of nopaline synthase had previously been inserted in the Asp718/BamHI and XbaI/PstI sites, respectively (38). Constructs consisting of a 339-bp fragment of the nopaline synthase promoter, gene-specific sequences in the sense orientation and a 253-bp fragment of the 3′ end of nopaline synthase were ligated to the PstI and HindIII sites of pCGN1547 and p35S∷A to make pNOS∷S and p35S∷A-NOS∷S, respectively. In p35S∷A-GUS-S, the β-glucuronidase (GUS) fragment containing nucleotides 787–1,809 was used as a linker between gene-specific fragments in the antisense and sense orientations. AG, CLV3, AP1, and PAN cDNA coding sequences used in this study contain nucleotides 301–855 (27), 3–291 (28), 445–854 (29), and 27–396 (30), respectively.
Agrobacterium-Mediated Transformation.
Agrobacterium strain ASE carrying DNA constructs in pCGN1547 was used to transform Arabidopsis plants (T0) by vacuum infiltration (39). Transformed Arabidopsis lines (T1) were selected on Murashige/Skoog (Sigma) plates containing kanamycin (50 μg/ml). Kanamycin-resistant seedlings were then transferred to soil. Phenotypic analysis of T1 and T2 plants is summarized in Table 1 and Table 2, respectively.
In Situ Hybridization.
The AG cDNA clone pCIT565 containing nucleotides 9–977 (27) was used to synthesize antisense and sense probes. 35S-labeled RNA probes were synthesized with Riboprobe in vitro Transcription Systems (Promega). The template was linearized with HindIII and transcribed by T7 RNA polymerase (antisense probe), or linearized with XhoI and transcribed by SP6 RNA polymerase (sense probe). Tissue was fixed in 1× PBS containing 4% paraformaldehyde/0.1% Triton X-100/0.1% Tween 20 at 4°C overnight. Fixed tissue was dehydrated with ethanol, cleared with xylene, embedded in paraffin (Paraplast Plus, Oxford Labware, St. Louis), and sectioned at 8 μm. In situ hybridization was performed as described by Drews et al. (40), with modifications by Sakai et al. (41). Exposure time was 8–10 days.
Western Blot Analysis.
Bud clusters (stages 1–12, including the inflorescence meristem) from one inflorescence were frozen and ground in liquid nitrogen, thawed in 30 μl of lysis buffer (50 mM Tris, pH 7.5/1 mM EDTA/100 mM NaCl/1% Nonidet P-40/0.1% SDS/0.1% Triton X-100/0.7% 2-mercaptoethanol/1 mM PMSF). The extract was mixed with 15 μl of 3× sample buffer (187 mM Tris, pH 6.8/6% SDS/30% glycerol/3% 2-mercaptoethanol/0.06% bromophenol blue), boiled for 5 min, and centrifuged (16,000 × g for 10 min at room temperature). Twenty microliters of the supernatant was separated on an SDS/12.5% polyacrylamide gel. The protein was transferred to nitrocellulose membrane (Schleicher & Schuell), probed with an AG-specific polyclonal antibody (42) and horseradish peroxidase-labeled secondary antibody (Amersham International), and detected with the enhanced chemiluminescence system (ECL; Amersham International).
To make constructs that produce dsRNA, gene-specific sequences in the antisense and sense configurations were either linked with the partial GUS gene and placed under control of the constitutive 35S promoter from cauliflower mosaic virus (p35S∷A-GUS-S), or controlled by the 35S promoter and the constitutive nopaline synthase promoter, respectively (p35S∷A-NOS∷S). A single RNA transcribed from the fusion gene in p35S∷A-GUS-S can potentially form a dsRNA stem with a single-stranded loop structure (Fig. 1). Genetic interference effects of sense, antisense, and dsRNAs corresponding to AG, CLV3, AP1, and PAN are outlined in Table 1. For each of these genes, p35S∷A-GUS-S constructs caused potent and specific genetic interference. However, p35S∷A-NOS∷S, p35S∷A and pNOS∷S constructs had either no, or weak, genetic interference effects. We will refer to transgenic plants carrying functional p35S∷A-GUS-S constructs by listing the gene name followed by RNAi. For unknown reasons, the sense construct corresponding to CLV3 caused toxicity in Agrobacterium and the sense construct of PAN resulted in very low transformation efficiency of crabs claw-1 (crc-1) plants. Therefore, interference effects of the CLV3 sense construct were not determined and only six crc-1 transgenic plants containing the PAN sense construct were analyzed.
AG dsRNA-Mediated Genetic Interference.
AG was chosen for initial characterization of RNAi in developing flowers. Arabidopsis flowers consist of four concentric whorls of organs. Most wild-type flowers have four sepals, four petals, six stamens, and two fused carpels, from the outermost first whorl to the innermost fourth whorl (Fig. 2A). Mutations in the AG gene cause homeotic alterations of the third and fourth whorls of organs in flowers (31). In severe ag loss-of-function mutants (Fig. 2B), the third whorl primordia develop into petals indistinguishable from those of the second whorl, and the fourth whorl develops into another ag flower, resulting in a repetitive pattern of sepals, petals, and petals (32).
The phenotypes produced by AG dsRNA are frequent and specific (Fig. 2 C–I). All but one of 236 transgenic plants showed ag mutant phenotypes. These AG (RNAi) mutants can be arranged into a phenotypic series based on the severity of the homeotic transformation in the third whorl and the extent of floral indeterminacy in the fourth whorl. Weak and intermediate AG (RNAi) mutant flowers showed partial homeotic transformation in the third whorl organs and slight floral indeterminacy (Fig. 2 E–H). However, intermediate/strong and strong AG (RNAi) mutant flowers showed complete transformation of the third whorl organs from stamens to petals and severe floral indeterminacy (Fig. 2 C, D, and I). Particularly, flowers from strong AG (RNAi) plants (Fig. 2C) are indistinguishable from those of strong ag mutant alleles such as ag-1 (Fig. 2B). Weak, intermediate, intermediate/strong, and strong AG (RNAi) mutants represent 16, 32, 43, and 9%, respectively, of the transgenic plant populations. In contrast, pNOS∷A-GUS-S, in which the nopaline synthase promoter was used to drive the fusion gene, and p35S∷A-NOS∷S constructs for AG caused very weak genetic interference in 2 out of 32 and 3 out of 124 transgenic plants, respectively (data not shown). Flowers from transgenic plants containing the AG antisense (n = 111) or sense (n = 95) construct are indistinguishable from those of wild-type plants (data not shown).
dsRNA Interferes with mRNA Accumulation.
In situ hybridization was performed to determine the target of dsRNA interference. The earliest expression of AG in wild-type flowers is in stage three, in those cells that will give rise to the third- and fourth-whorl organ primordia. Later, AG expression is restricted to the stamen and the carpel primordia (Fig. 3F) (27). The autoradiogram of the tissue hybridized with an AG anti-mRNA probe showed that hybridization signals declined with increasingly severe phenotypes in AG (RNAi) mutants (Fig. 3A–E), consistent with the observation that AG mRNA accumulation is decreased in AG (RNAi) mutants (Fig. 3G–J). These results suggest that endogenous mRNA is a target of dsRNA-mediated genetic interference. When used as a standard control for in situ hybridization, an AG sense probe hybridized with the tissue from AG (RNAi) mutants but not with that from wild type (Fig. 3K–N), suggesting that AG antisense RNA is produced in AG (RNAi) mutants. Reverse transcription–PCR analysis with GUS- and AG-specific primers also showed that expression levels of both strands of AG RNA from the fusion gene in p35S∷A-GUS-S increase with increasingly severe phenotypes (data not shown).
Furthermore, Western blot analysis of total protein from wild-type and AG (RNAi) mutants with an AG-specific polyclonal antibody (42) demonstrated that the severity of phenotypes is correlated with a reduction of the AG protein level in AG (RNAi) mutants. AG protein is weakly expressed in weak and intermediate AG (RNAi) mutants. In contrast, it is not detected at levels above background in intermediate/strong and strong AG (RNAi) mutants (Fig. 3O).
Genetic Interference by CLV3 and AP1 dsRNAs.
We further assessed the effectiveness and specificity of dsRNA with the CLV3 and AP1 genes. Plants with mutations in the CLV3 gene have enlarged meristems and extra floral organs, especially carpels (Fig. 4 C, F, and I) (33). The majority of CLV3 (RNAi) mutants (89%, n = 121) have flowers with extra carpels (Fig. 4H); however, only 2% of those plants (n = 108) also have extra sepals, petals, and stamens (Fig. 4E). In addition, some CLV3 (RNAi) mutant plants (26%, n = 121) have enlarged shoot apical meristems with distortions in phyllotaxy (Fig. 4B) and bifurcation, flattening, and broadening of the stem (data not shown). In contrast, only 1% (n = 176) of plants transformed with p35S∷A-NOS∷S for CLV3 have the extra carpel phenotype. clv3 mutant phenotypes were not observed in transgenic plants containing the CLV3 antisense construct (n = 273).
Mutations in the AP1 gene result in homeotic alterations of the outer two whorls and a partial or complete conversion of a floral meristem into an inflorescence meristem (34, 35). In weak ap1 mutant alleles, the first and second whorls consist of mosaic sepaloid organs and staminoid petals, respectively (Fig. 5B). In intermediate ap1 mutant alleles, flowers have leaf-like first whorl organs and leaf-like or staminoid second whorl organs (Fig. 5C). In strong ap1 mutant alleles, bract-like organs are produced in the first whorl and petals are usually absent in the second whorl (Fig. 5D). In addition, secondary flowers usually arise from the axils of the first whorl organs in flowers of intermediate and strong ap1 mutant alleles (Fig. 5 C and D). About 96% of transgenic plants (n = 260) containing the AP1 dsRNA-expressing construct, p35S∷A-GUS-S, produced flowers similar to those of ap1 mutant alleles (Fig. 5 E–H). Weak (Fig. 5E), intermediate (Fig. 5F), intermediate/strong (Fig. 5G), and strong (Fig. 5H) phenotypes were observed in 94, 1, 3, and 2%, respectively, of AP1 (RNAi) mutants (n = 249). In contrast, transgenic plants containing the AP1 construct in the antisense orientation (6%, n = 140) had very weak mutant phenotypes (data not shown). The AP1 sense construct did not cause mutant phenotypes in transgenic plants (n = 62; data not shown).
One CLV3 (RNAi) T1 plant and 6 AP1 (RNAi) T1 plants of variable severity were selfed to examine the inheritance of genetic interference (Table 2). The progeny from each selected plant showed similar severity of phenotypes to those of the selfed parents, and the severity of phenotypes is constant between mutant siblings of each lineage. In addition, the progeny of AP1 (RNAi) plants had 3:1 (mutant/wild type) segregation ratios, suggesting that each of the 6 AP1 (RNAi) T1 plants contained one copy of the transgene. This result indicates that dsRNA-expressing constructs, which are integrated into the plant genome, are stably inherited in a Mendelian fashion, and that the RNAi effect persists to, or recurs in, new generations of plants.
RNA-Mediated Interference with PAN.
Flowers of plants mutant for pan are characterized by an increase in the organ number in the first two whorls, and a decrease in the organ number in the third whorl. Mutant flowers usually have five sepals, five petals, five stamens, and two carpels (36). When introduced into wild-type plants, the dsRNA-expressing construct of PAN caused either no, or weak, extra organ phenotypes. Reverse transcription–PCR analysis showed that PAN mRNA was reduced by 30–90% in PAN (RNAi) plants compared with wild-type plants (data not shown), suggesting that a small portion of PAN activity is sufficient for its function in wild-type plants. This hypothesis is consistent with results from previous immunohistochemical analysis showing that the mutant pan-1 and pan-2 alleles, with high expressivity of the extra organ phenotype, are likely null alleles (30).
Whereas flowers homozygous for strong mutant alleles show high penetrance of the extra organ phenotype, only the first few flowers from homozygotes of weak mutant alleles show the phenotype. However, both strong and weak pan alleles cause high penetrance of unfused carpel phenotypes in a crc-1 genetic background (Fig. 6 A, B, D, and E) (30, 44), suggesting that crc mutants provide a more sensitive background than wild type in which to observe phenotypic effects of PAN reduction-of-function mutations. Therefore, RNA-expressing constructs corresponding to PAN were introduced into a crc-1 homozygous background to further assess the potential of RNA-mediated interference with PAN. Similar to pan alleles, PAN dsRNA-expressing constructs, p35S∷A-GUS-S (87%, n = 126) and p35S∷A-NOS∷S (27%, n = 66), caused extra organ number and unfused carpels in crc-1 (Fig. 6 C and F). Antisense (55%, n = 76) and sense (33%, n = 6) sequences corresponding to PAN have similar RNAi effects as well (data not shown), suggesting that low levels of dsRNAs might be produced in such a case and weak interference with PAN activity is sufficient to confer an unfused carpel phenotype in crc-1.
This study shows that dsRNA-mediated genetic interference can operate in A. thaliana to efficiently induce sequence-specific inhibition of gene function. Although the technique of RNA microinjection has been widely used in C. elegans (13–15, 44), Drosophila (20, 21), and planarians (23), methods for RNA injection into zygotes of A. thaliana are not available. However, Agrobacterium-mediated transformation provides a convenient and efficient method to introduce dsRNA-expressing constructs into the plant genome. Therefore, RNAi in transgenic plants is persistent and inherited instead of being transient and unstable as in RNA-injected animals (13–15, 20, 21, 23, 45) and transiently transfected cells (22). In addition, inducible and tissue-specific promoters might be used to obtain regulated RNAi.
In this study, two kinds of dsRNA-expressing constructs, p35S∷A-GUS-S and p35S∷A-NOS∷S, were used to investigate RNAi effects. p35S∷A-NOS∷S is less potent at inducing genetic interference than p35S∷A-GUS-S, perhaps because of unequal expression levels of sense and antisense RNAs by two promoters of different strength. The nopaline synthase promoter is much weaker than the 35S promoter, suggested by the observation that pNOS∷A-GUS-S has weaker genetic interference than p35S∷A-GUS-S. These results suggest that equal and high levels of both strands of each RNA in each cell are essential for inducing potent RNAi. If this is true, use of two strong promoters of similar strength should improve RNAi effects of dsRNA-expressing constructs in which sense and antisense RNAs are produced separately; however, use of two identical promoters in a construct should be avoided to prevent cosuppression (46).
dsRNAs corresponding to four genes selected in this study caused potent and specific genetic interference, suggesting that dsRNA-mediated gene silencing can occur in the tissues where these genes normally function. In addition, a phenotypic series can be obtained from RNAi mutants. The fact that the severity of phenotypes varied between T1 individuals is possibly because of variable transgene copy number and/or positional effects of particular DNA insertion events. However, our results suggest that severity of phenotypes in AP1 (RNAi) T1 plants is not related to the transgene copy number.
CLV3 dsRNA seems predominantly to block gene function in a subset of the cells where it is normally expressed. CLV3 is expressed in the inflorescence and the floral meristems (28). Mutations in the CLV3 gene cause enlarged meristems and extra floral organs (33). About 89% of CLV3 (RNAi) plants have flowers with extra carpels but only 26% of CLV3 (RNAi) plants have enlarged inflorescence meristems. This result suggests a strong suppression of the CLV3 gene function in the center of the floral meristem but less suppression of its function in the inflorescence meristem. It is probably because of differential activity of the 35S promoter which drives expression of dsRNA in these tissues. It is also possible that some tissues could partially resist RNAi (25), or that some phenotypes (such as enlarged inflorescence meristems) are less sensitive to the level of gene activity.
When used as controls for RNAi experiments, the sense and antisense constructs of PAN also had the ability to induce genetic interference in a crc-1 homozygous background; so did the AP1 antisense construct in wild-type plants. It has been suggested that low levels of dsRNA might be produced from transgenes that are designed to produce only antisense or only sense RNA, via the readthrough transcription from transgenes arranged as an inverted repeat, or transcription from a transgene whose 3′ end is adjacent to an endogenous promoter (19, 24, 25, 47). Alternatively, it seems possible that cellular RNA-dependent RNA polymerase could be involved in the conversion of single-stranded RNA to dsRNA in a cell-specific manner, suggested by the cloning and in vitro catalytic analysis of an RNA-dependent RNA polymerase from tomato (25, 48).
In situ hybridization revealed a correlation between decreasing levels of AG mRNA accumulation and increasing severity of phenotypes in AG (RNAi) plants, suggesting that the mechanism blocking mRNA accumulation could be responsible for dsRNA interference. This result is consistent with previous findings that endogenous mRNA is a target of dsRNA-mediated genetic interference (13, 14, 22, 23). In addition, a recent report of isolation of an RNaseD homolog from C. elegans mutants which are resistant to RNAi suggests that RNAi works by dsRNA-directed, enzymatic RNA degradation (49).
Whatever the mechanism by which RNAi acts to reduce specific mRNA levels, the experiments described here show that it is a useful method for determining the loss-of-function phenotypes of genes involved in development and meristem activity in A. thaliana.
We thank John Bowman and Yuval Eshed for critical reading of the manuscript; Xavier Ambroggio, Catherine Baker, Chieh Chang, Toshiro Ito, Jeff Long, Kazuaki Ohashi, Carolyn Ohno, G. Venugopala Reddy, Doris Wagner, Frank Wellmer, and Eva Ziegelhoffer for their comments on the manuscript; Amani Zewail for technical assistance; and Toshiro Ito for providing the anti-AG antibody. This work was supported by National Institutes of Health Grant GM45697 to E.M.M.
↵* To whom reprint requests should be addressed. E-mail: meyerow{at}cco.caltech.edu.
Article published online before print: Proc. Natl. Acad. Sci. USA, 10.1073/pnas.060034297.
Article and publication date are at www.pnas.org/cgi/doi/10.1073/pnas.060034297
- dsRNA,
- double-stranded RNA;
- RNAi,
- RNA interference;
- β-glucuronidase
- Accepted January 24, 2000.
- Copyright © The National Academy of Sciences
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