[NEW] Diversification of gene function: homologs of the floral regulator FLO/LFY control the first zygotic cell division in the moss Physcomitrella patens | lfy – Vietnamnhanvan

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After fertilization, the zygote undergoes dynamic changes in chromosomal and cytoplasmic organization, and begins the cell cycles that eventually lead to formation of the multicellular embryo. Specific transcription factors that initiate this cascade of events in land plants have not been identified. We have identified two FLO/LFY genes, PpLFY1 and PpLFY2, that regulate the first cell division after formation of the zygote in the moss Physcomitrella patens. The deduced amino acid sequences of the two PpLFY genes are 94.8% identical to each other and show similar expression patterns. While fertilization occurred in the PpLFY disruptants, the development of double disruptant zygotes was arrested at the single-cell stage. When the double disruptants, as the female parent, were crossed with the wild type, as the male parent, normal sporophytes were formed, supporting the notion that the PpLFY genes function after fertilization to regulate the first mitotic cell division in zygotes. The rare sporophytes that formed on the PpLFY double disruptants showed mostly normal organogenesis, but had abnormalities in the pattern of cell division, supporting a role of PpLFY genes in regulating cell division. The FLO/LFY genes in angiosperms are conserved master regulators of floral identity without any obvious effects on cell division. By contrast, our study suggests that FLO/LFY genes have functions throughout sporophyte development in the basal land plant lineages.

We report the characterization of two FLO/LFY genes in P. patens, and show that the FLO/LFY genes play a key role in the progression of the first cell division of the zygote. In addition, they are important for later sporophyte development.

The moss Physcomitrella patens has been recognized as a useful model for the study of plant embryogenesis and development( Cove et al., 1997 ; Sakakibara et al., 2003 ) for several reasons. First, the exceptionally high efficiency of homologous recombination in P. patens makes it the only land plant in which gene targeting is feasible ( Schaefer and Zrÿd, 1997 ). Second, the eggs, zygotes and young sporophytes of P. patens are located solely in the cavity of its egg-bearing organ, the archegonium, and are thus easily accessible for observation, while those in seed plants are relatively difficult to observe. Third, P. patens has a multicellular autotrophic body in its haploid generation,permitting maintenance of mutants with severe defects in their diploid generation.

Multicellular organisms begin their diploid development from a single-cell stage: the zygote. The proper initiation of mitotic cell cycles in the fertilized egg is a key to the progression of development in both land plants and metazoans. In land plants, however, the regulatory mechanisms of this critical step remain mostly unknown. The first cell division of the zygote in land plants is usually asymmetric, forming a cytoplasm-rich apical cell and a vacuole-rich basal cell, although the plane of cell division and the orientation of the apical-basal axis to the principal axis of the egg-bearing organ vary among plant lineages ( Wardlaw,1955 ). Among land plants, only one gene, GNOM, which functions in vesicle trafficking, has been characterized to be involved in apical-basal polarity and in the normal division of the zygote( Mayer et al., 1993 ; Geldner et al., 2003 ). However, transcription factors affecting the first cell division of zygotes have not been identified in land plants.

Physcomitrella patens Bruch & Schimp subsp. patens( Ashton and Cove, 1977 ) was cultured as described previously( Nishiyama et al., 2000 ). To observe the gametangia and sporophytes, protonemata that had been vegetatively propagated on 9 cm Petri dishes with BCDAT medium( Nishiyama et al., 2000 ) were inoculated onto sterile 42-mm diameter peat pellets (Jiffy-7; Jiffy Products International AS, Kristansand, Norway) using tweezers. The cylindrical pellets had been prepared by immersion in distilled water to permit them to expand,and then autoclaved in a plastic box. Distilled water was then poured to just below the upper surface of the expanded cylindrical pellets. The mosses inoculated on the peat pellets were cultured at 25°C under 16-hour light/8-hour dark conditions for 1 to 1.5 months, and then transferred to 15°C under 8-hour light/16-hour dark conditions for induction of gametangia development. Under these conditions, first antheridia and then archegonia differentiated at the shoot apices of the gametophores. Because of the wet conditions, fertilization occurred without any additional treatment. Sporophytes were visible under stereomicroscope 4 weeks after the transfer to 15°C and were collected after an additional month of culture for experiments using spores. To estimate the germination rate, the spores were spread on solidified BCDAT medium, and the germinated protonemata were counted after 1 week.

Genomic structures of PpLFY1 and PpLFY2 in the wild type and the targeted lines. The gene structures of the wild-type PpLFY1(A) and PpLFY2 (B). Restriction enzyme recognition sites are indicated on each gene: b, BglII; e, EcoRI. In A,B,D-G the boxes indicate exons and the lines between the boxes, introns; circles,putative start codons and squares, putative stop codons. The DNA fragments that were used as probes in the Southern analyses are indicated with thick lines below each gene in A and B. (C) Southern analyses of PpLFY1 and PpLFY2 in the wild type. The restriction enzymes, probes, and stringency used are indicated above or below each image. (D-G) Genomic structures of PpLFY1 (D,F) and PpLFY2 (E,G) in PpLFY1-GUS lines (D), PpLFY2-GUS lines (E), PpLFY1-dis and PpLFY1-PpLFY2-dis lines (F),PpLFY2-dis and PpLFY1-PpLFY2-dis lines (G). uidA, uidA coding region;nos-ter, nopaline synthase polyadenylation signal; NPTII, NPTII expression cassette; HPT, HPT expression cassette. The sense directions of uidA,nptII, and hpt are indicated with arrows. The scale bar applies to A,B,D-G.

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Genomic structures of PpLFY1 and PpLFY2 in the wild type and the targeted lines. The gene structures of the wild-type PpLFY1(A) and PpLFY2 (B). Restriction enzyme recognition sites are indicated on each gene: b, BglII; e, EcoRI. In A,B,D-G the boxes indicate exons and the lines between the boxes, introns; circles,putative start codons and squares, putative stop codons. The DNA fragments that were used as probes in the Southern analyses are indicated with thick lines below each gene in A and B. (C) Southern analyses of PpLFY1 and PpLFY2 in the wild type. The restriction enzymes, probes, and stringency used are indicated above or below each image. (D-G) Genomic structures of PpLFY1 (D,F) and PpLFY2 (E,G) in PpLFY1-GUS lines (D), PpLFY2-GUS lines (E), PpLFY1-dis and PpLFY1-PpLFY2-dis lines (F),PpLFY2-dis and PpLFY1-PpLFY2-dis lines (G). uidA, uidA coding region;nos-ter, nopaline synthase polyadenylation signal; NPTII, NPTII expression cassette; HPT, HPT expression cassette. The sense directions of uidA,nptII, and hpt are indicated with arrows. The scale bar applies to A,B,D-G.

Genomic DNA was extracted from a mixture of protonemata and gametophores of P. patens by the CTAB (Hexadecyltrimethyl-ammonium bromide) method( Murray and Thompson, 1980 ). The blotting and hybridization of 1 μg of digested genomic DNA were performed as described previously ( Shindo et al., 2001 ), using the gPpLFY1 and gPpLFY2 DNA fragments as probes ( Fig. 1A,B ).

Total RNA was extracted from protonemata, gametophores, or sporophytes using RNeasy plant mini-kit (Qiagen, Hilden, Germany). The protonemata were grown on solidified BCDAT medium for 6 days. The gametophores were grown on the solidified BCD medium ( Nishiyama et al., 2000 ) for 80 days. Various developmental stages of sporophytes, excluding archegonium tissue, were collected from mosses grown on peat pellets for a month after the transition to 15°C. The cDNAs were synthesized from 1 μg of total RNA from each tissue using SuperscriptII reverse transcriptase (Invitrogen, Carlsbad, CA, USA) and oligo(dT) 16 primer according to the manufacturer’s instructions. PCR was performed with TaKaRa Ex Taq Hot Start Version (Takara Bio Inc., Otsu, Japan), using the following gene-specific primers: for PpLFY1, pL1-NNsense1(5′-CGGTGGCGGAAGGTAGCAAAAGTGAAAGCA-3′) and L1e2spe(5′-CCTCCTCCAAGAATTTCCCACATTGCTCG-3′), and for PpLFY2,PpLFY2-5′ (5′-GCTCGGCTGCATTCCAGCTGCACTCTC-3′) and L2e2-5-2(5′-AATGTGCTGCACCTCTTCCAGAAACTTTC-3′). The PCR products were sequenced directly to confirm the amplification of the expected fragments.

Based on the cDNA sequences of PpLFY1 and PpLFY2(Himi et al., 2001), the 5′ and 3′ genomic regions of these cDNAs were amplified using TAIL-PCR (Liu and Whittier,1995) and sequenced. These cDNA and genomic DNA sequences were then used to construct targeting plasmids to make PpLFY1-GUS, PpLFY2-GUS,PpFLY1-dis, PpLFY2-dis and PpLFY1-PpLFY2-dis lines in addition to PpLFY ectopic expression lines in A. thaliana.

Genomic sequences of PpLFY1 and PpLFY2 were used to design and synthesize primer sets for the amplification of the majority of both PpLFY1 (2871 bp) and PpLFY2 (2852 bp) genomic DNA:L1-1int1 (5′-CTTGGCTTTAGACTCCTCTCATCGGTCG-3′) and L1-3U1(5′-TTGGTCAAGATCTCATGGCCGCAGCTG-3′), and PpLFY2-5′(5′-GCTCGGCTGCATTCCAGCTGCACTCTC-3′) and PpLFY2-3′(5′-CTACAAAATAGTGTACAAGGGCTCATTCG-3′). Each amplified DNA fragment was cloned into the pGEM-T Easy vector (Promega, Madison, WI, USA), thereby generating pgPpLFY1 and pgPpLFY2.

A partial 1367 bp fragment of PpLFY1 genomic DNA ranging from the second exon to just before the stop codon was also cloned into the pGEM-T vector (Promega). A SalI-SmaI fragment was excised from this plasmid and inserted into a SalI-ClaI site of the pGUS-NPTII-1 plasmid that contained the coding sequence of uidA(Jefferson et al., 1987), the nopaline synthase polyadenylation signal (nos-ter), and an NPTII cassette(Nishiyama et al., 2000),thereby creating an in-frame fusion of the PpLFY1 and uidAgenes. A 943 bp DNA fragment containing the 3′ region of PpLFY1gene, produced by TAIL-PCR using the gene-specific primer PpLFY-LA1(5′-CGGATCTGGTATGTCCCCACGAAACTCC-3′), was inserted into the SmaI site of this plasmid. The resulting plasmid was linearized with BamHI and SalI for generation of the PpLFY1-GUS lines. Of 78 independent stable transformants, 18 were identified by PCR to have the fusion construct in the PpLFY1 locus. Southern analyses of 12 of the 18 candidate lines showed that three lines (PpLFY1-GUS-1, 2 and 3) contained a single insertion in the PpLFY1 locus (see Fig. S1A,C in the supplementary material), while the other nine lines tested contained multiple copies (data not shown).

A 1460 bp fragment containing a partial sequence of the 5′ region of the PpLFY2 genomic DNA was amplified using the PpLFY2-5′ and L2a-Hi3 (5′-CAAGCTTCGGCTTTCTCTGCCGGCCAGCGC-3′) primers. A 1398 bp fragment of the 3′ region of the PpLFY2 genomic DNA was amplified using the L2s-Ba1 (5′-CGGATCCGAAGGTTGAAGGAGCTATTCAAG-3′)and PpLFY2-3′ primers. These fragments were inserted into pGUS-NPTII-1,creating an in-frame fusion of the PpLFY2 and uidA genes. The resulting plasmid was linearized with NotI and SalI and used to generate the PpLFY2-GUS lines. Of 80 independent stable transformants,44 were identified by PCR to have the fusion construct in the PpLFY2locus. Southern analyses of 15 of the 44 candidate lines showed that 11 lines(PpLFY2-GUS-1 to -11) contained a single insertion in the PpLFY2locus (see Fig. S1B,D in the supplementary material; data not shown), while the other lines tested contained multiple copies (data not shown).

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To disrupt the PpLFY1 gene, the 5′ SalI-HindIII fragment and the 3′ XbaI fragment of pgPpLFY1 were inserted into the SalI-HindIII and XbaI sites of pHTS14, respectively. pHTS14 contains an HPT cassette composed of the CaMV 35S promoter, the hygromycin B phosphotransferase(hpt) gene, and the nos-ter sequence between multiple cloning sites. The resulting plasmid was used for transformation after linearization with NotI and then used to generate the PpLFY1-dis and PpLFY1-PpLFY2-dis lines. To generate PpLFY1-dis lines, 17 of 38 lines randomly selected from 81 independent stable transformants were identified by PCR as having a disrupted PpLFY1 locus (data not shown). Southern analyses of 10 of the 17 candidate lines showed that 8 lines (PpLFY1-dis-1 to 8) contained a single insertion in the PpLFY1 locus (see Fig. S2A,C in the supplementary material), while the other lines contained multiple copies (data not shown).

To disrupt the PpLFY2 gene, the fragment of PpLFY2between the HindIII sites located in the second intron and the third exon of pgPpLFY2 was replaced with an NPTII cassette. The resulting plasmid was used for transformation after linearization with EcoRI and NotI. To generate the PpLFY2-dis line, 5 of 56 lines randomly selected from 79 independent stable transformants were identified by PCR as having a disrupted PpLFY2 locus (data not shown). Southern analyses of the 5 candidate lines showed that a single line (PpLFY2-dis-1) contained a single insertion in the PpLFY2 locus (see Fig. S2B,D in the supplementary material), while the other lines contained multiple copies (data not shown).

To generate double disruptants, an HPT cassette was inserted into the third exon of PpLFY1 in the PpLFY2-dis-1 line. Of 47 independent stable transformants, 20 were identified by PCR as having a disrupted PpLFY1locus (data not shown). Southern analyses of 18 of the 20 candidate lines showed that 6 lines (PpLFY1-PpLFY2-dis-1 to -6) contained a single insertion in the PpLFY1 locus, in addition to the disrupted PpLFY2locus (see Fig. S2D,E in the supplementary material), while the other lines contained multiple copies (data not shown).

[NEW] Last Fiscal Year (LFY) Definition | lfy – Vietnamnhanvan

What Is a Last Fiscal Year (LFY)?

The term last fiscal year (LFY) refers to the most recent 12-month accounting period that a business uses to determine its annual financial performance. The Securities and Exchange Commission (SEC) requires businesses to list their last fiscal year’s revenue, in addition to other financial figures measured on a fiscal year basis. Analysts, investors, and corporate management often use figures and metrics from a company’s last fiscal year to make forecasts about its financial performance.

Key Takeaways

  • A last fiscal year is the most recent 12-month accounting period.
  • The LFY is used by a company to determine its annual financial performance.
  • The SEC requires companies to include information from their last fiscal year on financial statements and reports, including their annual filings.
  • Investors and analysts can use a company’s LFY to make predictions about its future performance.
  • One-time non-operating events should be noted as such because they may skew a company’s metrics.

Understanding Last Fiscal Years (LFYs)

A fiscal year is an annual period generally used by companies to report their financial statements for accounting purposes. The term fiscal year is also referred to as a budget year. Governments also use operate through fiscal years and report financial data once that period is up. Fiscal years run for 12 full months and are characterized by the year-end.

A company’s fiscal year may not be the same as a calendar year. This means they don’t necessarily run from January to December. Some fiscal years run for the 12-month period between July 1 and June 30. Others may have their fiscal year between Oct. 1 and Sept. 30 of each year. Corporations choose the 12-month period for which they report their financial statements based on the type of company and the seasonality of the business.

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The most current 12-month period reported by a company is called its last fiscal year. Financial information is submitted on a timely basis and at the fiscal year-end. The LFY is used as a way to determine a company’s financial performance. As mentioned above, the SEC requires companies to include information from their last fiscal year on a number of their financial statements and reports, including their 10-K and 10-Q filings.

Information reported by companies in their last fiscal years provides a lot of valuable information for investors and financial professionals. For instance, an analyst can use information from the last fiscal period to make forecasts about the future of different companies. They can try to predict whether or not a business’s current performance will outdo that of the previous fiscal year.

A fiscal year-end normally occurs at the end of a quarter.

Special Considerations

While there is a chance that the last fiscal year may help predict future performance, there are exceptions to this rule. For instance, the inclusion of one-time financial anomalies in the last fiscal year’s results may cause an ineffective comparison. That’s because one-time non-operating events could skew a company’s metrics.

Let’s say ABC Corporation sold a factory for $1 million and reported the cash as revenue in the last fiscal year’s financial statements. Conducting normal analysis and including this information from the LFY will provide inaccurate results. Unless it is specified that this extra sum of money didn’t originate from its regular day-to-day operations, individuals may mistakenly believe that the company’s operations generated an extra million dollars and may continue the trend going forward.

Example of a Last Fiscal Year

Let’s use the hypothetical example of ABC Corporation from the section above to show how LFYs work. Suppose the company’s fiscal year starts on April 1 every year and ends on March 31, and it is currently July. If it were to list its revenue from the last fiscal year, it would show the results that took place from April 1 of the previous year to March 31 of the current year.


VP vs LFY TI7 Semi Final Highlights The International 2017 Dota 2


VP vs LFY TI7 Semi Final Highlights The International 2017 Dota 2

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VP vs LFY TI7 Semi Final Highlights The International 2017 Dota 2

Liquid vs LFY TI7 [EPIC] LB Final Highlights The International 2017 Dota 2


Liquid vs LFY TI7 [EPIC] LB Final Highlights The International 2017 Dota 2

Liquid vs LFY TI7 [EPIC] LB Final Highlights The International 2017 Dota 2

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LFY vs VP Game 2 | Dota 2 TI7 2017 | LGD FY vs VIRTUS PRO

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