Time of bloom

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By Austin Mudd - Spring 2013


To Do

  • Write introduction and procedure sections
  • Continue searching for grape orthologs


Flowering Introduction

The factors affecting time of flowering from the Max Planck Institute for Plant Breeding Research

Fornara et al., 2010

"Plants initiate flowering after a period of vegetative development. During this process, called floral induction, the shoot apical meristem starts to produce flowers instead of leaves. The timing of floral induction is controlled by sophisticated regulatory networks that monitor changes in the environment, ensuring that flowering occurs under conditions most likely to maximize reproductive success and seed production. In the model plant species Arabidopsis thaliana ?180 genes have been implicated in flowering-time control based on isolation of loss-of-function mutations or analysis of transgenic plants. This SnapShot presents a subset of these genes and proteins, each organized according to its spatial activity in the leaves or the shoot apical meristem of the plant. Strikingly, several genes act more than once and in several tissues during floral induction. Many of these genes occur in a network of six major pathways: the photoperiod and vernalization pathways control flowering in response to seasonal changes in day length and temperature; the ambient temperature pathway responds to daily growth temperatures; and the age, autonomous, and gibberellin pathways act more independently of environmental stimuli. The six pathways converge to regulate a small number of “floral integrator genes,” encoded by different classes of proteins, which govern flowering time by merging signals from multiple pathways. These integrator genes include FLOWERING LOCUS T (FT) and SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1), which both rapidly promote floral development. In addition, responses to other environmental stimuli, such as the balance of different wavelengths of light or nutrient availability, also influence flowering time, but how these processes interact with the pathways described here is not fully understood."


Higgins et al., 2010

"The switch from vegetative growth (the production of stems and leaves) to reproductive growth (the production of flowers) is an important developmental step in the life cycle of plants. Flowering needs to occur when conditions for pollination and seed development are optimal and consequently most plants restrict flowering to a specific time of year. They commonly achieve this by using reliable environmental cues such as day length (photoperiod) and temperature. In addition, nutrient and water availability and plant size can be important.
The genes and molecular mechanisms controlling flowering have been extensively studied in the model dicot Arabidopsis thaliana, subsequently Arabidopsis ... As part of this study the Arabidopsis flowering pathways were curated in Arabidopsis Reactome ... to provide an electronic knowledge resource allowing for further developments such as integration with protein-protein interaction datasets, overlaying with microarray data and electronic projection into all newly sequenced plant genomes. Using this we compiled a list of genes and gene families with a known role in flowering time in Arabidopsis.
Flowering time has also been extensively studied in crop species ... Flowering time is important for adaptation to specific environments and the world's major crop species provide a particularly interesting opportunity for study because they are grown in areas outside the ecogeographical limits of their wild ancestors. In addition, they are adapted to different farming practices such as fall (autumn) sowing or spring sowing in temperate regions. Adaptation to different environments and practices has been achieved by manipulation of flowering time responses and this makes flowering pathways an excellent system for comparison between and within domestic and wild species."


Jung et al., 2012

"Plants switch to the reproductive phase of development when environmental and endogenous factors are the most favourable for reproductive success and seed production. This proper timing is the result of elaborate regulatory networks that coordinate the external stimuli with endogenous cues, inducing the expression of genes that initiate the floral transition at the shoot apical meristem (SAM).
Much of our current understanding of the floral initiation process is derived from studies using Arabidopsis thaliana as the model system. More than 180 Arabidopsis genes have been identified that play a role in regulating flowering time, and these genes have been organised into six major pathways ... Although the photoperiod and vernalisation pathways monitor seasonal changes in day length or temperature and, hence, initiate flowering in response to exposure to long days or prolonged cold temperatures, the ambient temperature pathway coordinates the response to daily growth temperatures. The autonomous pathway together with those involving age or gibberellin constitutes the rest of the floral pathways, which function more independently of external stimuli. These pathways are integrated by downstream target genes including LEAFY (LFY), FLOWERING LOCUS T (FT) and SUPPRESSOR OF CONSTANS1 (SOC1), with their resulting outcomes conveyed to floral meristem identity genes such as APETALA1 (AP1) at the SAM that triggers the flowering process ...
Flowering is one of the most important agronomic traits influencing crop yield. There is thus a great necessity for research that examines the molecular control of this fundamental process in important crop species. This knowledge is critical for the breeding of climate change resilient crop varieties. Soybean, a major food crop, is also a member of the large and diverse legume family, which has the unique capability of forming nitrogen-fixing symbioses with soil microorganisms and has thus been used as part of sustainable agricultural practices for thousands of years. Soybean is distributed broadly across latitudes and is cultivated as different maturity groups, with each having a narrow range of latitudinal adaptation. Unlike Arabidopsis, soybean can undergo a reversion of flowering when plants are shifted from flowering inductive to non-inductive conditions ... In addition, soybean also follows a floral developmental plan that is distinct from that of Arabidopsis ... Therefore, an understanding of the molecular mechanisms underlying these soybean traits is of fundamental and practical interest."


  • General plant anatomy / process of flowering
  • Why flowering is important to study / general impact


Light Signaling Pathway

Jung et al., 2012

"Light is one of the main environmental regulators of flowering in plants. Plants sense the time of day and season of year by monitoring the light environment through light signalling pathways ... In Arabidopsis, photoperiod pathway genes together with photoreceptor genes and circadian clock components take part in light signalling pathways. The number of known Arabidopsis flowering genes involved in these pathways is 48 ... The key Arabidopsis genes involved in the light signalling pathway include the CONSTANS (CO), PHYTOCHROME (PHY) and CRYPTOCHROME (CRY), CIRCADIAN CLOCK ASSOCIATED 1 (CCA1), LATE ELONGATED HYPOCOTYL (LHY) and PSEUDO-RESPONSE REGULATOR 1 [PRR1, also called TIMING OF CAB EXPRESSION 1 (TOC1)] genes."


Vernalization Pathway

Jung et al., 2012

"Vernalisation involves plants that require prolonged periods of low temperature to initiate flowering. The vernalisation pathway in Arabidopsis involves 32 genes".


Autonomous Pathway

Jung et al., 2012

"Autonomous pathways in plants are activated in response to endogenous changes that are independent from the environmental cues leading to flowering ... There are 17 genes ... involved in the Arabidopsis autonomous pathway".


Ambient Temperature Pathway

Jung et al., 2012

"Plants respond to ambient temperature changes to modulate their flowering times ... The ambient temperature pathway in Arabidopsis involves 16 genes".


Procedure

  • Overall procedure for finding the genes, running with GenSAS, determining SSRs, etc


Gallery of Arabidopsis Flowering Pathways


105 Flowering Genes

This table lists all of the genes involved in the ambient temperature, autonomous, light signaling, and vernalization pathways. All Arabidopsis genes are compiled from Jung et al., 2012. All potential orthologs are found via UniProt Grape or UniProt Strawberry nomenclature search.

Arabidopsis Locus Other Names AA Sequence Pathway Potential Ortholog
AT1G01060 LATE ELONGATED HYPOCOTYL, LATE ELONGATED HYPOCOTYL 1, LHY, LHY1 TAIR Light signaling
AT1G02580 EMB173, EMBRYO DEFECTIVE 173, FERTILIZATION INDEPENDENT SEED 1, FIS1, MEA, MEDEA, SDG5, SET DOMAIN-CONTAINING PROTEIN 5 TAIR Autonomous, Vernalization
AT1G04400 AT-PHH1, ATCRY2, CRY2, CRYPTOCHROME 2, FHA, PHH1 TAIR Light signaling
AT1G09570 ELONGATED HYPOCOTYL 8, FAR RED ELONGATED 1, FAR RED ELONGATED HYPOCOTYL 2, FHY2, FRE1, HY8, PHYA, PHYTOCHROME A TAIR Light signaling Grape
AT1G13260 EDF4, ETHYLENE RESPONSE DNA BINDING FACTOR 4, RAV1, RELATED TO ABI3/VP1 1 TAIR Light signaling
AT1G20330 COTYLEDON VASCULAR PATTERN 1, CVP1, FRILL1, FRL1, SMT2, STEROL METHYLTRANSFERASE 2 TAIR Vernalization
AT1G22770 FB, GI, GIGANTEA TAIR Light signaling
AT1G25560 EDF1, ETHYLENE RESPONSE DNA BINDING FACTOR 1, TEM1, TEMPRANILLO 1 TAIR Light signaling
AT1G26790 TAIR Light signaling
AT1G29160 TAIR Light signaling
AT1G30970 SUF4, SUPPRESSOR OF FRIGIDA4 TAIR Vernalization
AT1G31814 FRIGIDA LIKE 2, FRL2 TAIR Vernalization
AT1G47250 20S PROTEASOME ALPHA SUBUNIT F2, PAF2 TAIR Vernalization
AT1G53090 SPA1-RELATED 4, SPA4 TAIR Light signaling
AT1G62830 ARABIDOPSIS LYSINE-SPECIFIC HISTONE DEMETHYLASE, ATLSD1, ATSWP1, LDL1, LSD1, LSD1-LIKE 1, LYSINE-SPECIFIC HISTONE DEMETHYLASE, SWP1 TAIR Ambient temperature, Autonomous
AT1G65480 FLOWERING LOCUS T, FT TAIR Ambient temperature, Flowering integrator Grape, Strawberry
AT1G68050 "FLAVIN-BINDING, KELCH REPEAT, F BOX 1", ADO3, FKF1 TAIR Light signaling
AT1G68840 ATRAV2, EDF2, ETHYLENE RESPONSE DNA BINDING FACTOR 2, RAP2.8, RAV2, RELATED TO ABI3/VP1 2, RELATED TO AP2 8, TEM2, TEMPRANILLO 2 TAIR Light signaling
AT1G77080 AGAMOUS-LIKE 27, AGL27, FLM, FLOWERING LOCUS M, MADS AFFECTING FLOWERING 1, MAF1 TAIR Ambient temperature, Flowering integrator, Vernalization
AT1G77300 ASH1 HOMOLOG 2, ASHH2, CAROTENOID CHLOROPLAST REGULATORY1, CCR1, EARLY FLOWERING IN SHORT DAYS, EFS, LAZ2, LAZARUS 2, SDG8, SET DOMAIN GROUP 8 TAIR Vernalization
AT2G06255 ELF4-L3, ELF4-LIKE 3 TAIR Light signaling
AT2G16780 MSI2, NFC2, NUCLEOSOME/CHROMATIN ASSEMBLY FACTOR GROUP C 2 TAIR Autonomous, Vernalization
AT2G17770 ATBZIP27, BASIC REGION/LEUCINE ZIPPER MOTIF 27, BZIP27, FD PARALOG, FDP TAIR Ambient temperature, Meristem identity
AT2G18790 HY3, OOP1, OUT OF PHASE 1, PHYB, PHYTOCHROME B TAIR Light signaling Grape
AT2G18870 VEL3, VERNALIZATION5/VIN3-LIKE 3, VIL4, VIN3-LIKE 4 TAIR Autonomous
AT2G18880 VEL2, VERNALIZATION5/VIN3-LIKE 2, VIL3, VIN3-LIKE 3 TAIR Autonomous
AT2G18915 ADAGIO 2, ADO2, LKP2, LOV KELCH PROTEIN 2 TAIR Light signaling
AT2G19520 ACG1, ATMSI4, FVE, MSI4, MULTICOPY SUPPRESSOR OF IRA1 4, NFC04, NFC4 TAIR Ambient temperature, Autonomous
AT2G22540 AGAMOUS-LIKE 22, AGL22, SHORT VEGETATIVE PHASE, SVP TAIR Ambient temperature, Vernalization Grape
AT2G23380 CLF, CURLY LEAF, ICU1, INCURVATA 1, SDG1, SET1, SETDOMAIN 1, SETDOMAIN GROUP 1 TAIR Autonomous, Vernalization
AT2G25930 EARLY FLOWERING 3, ELF3, PYK20 TAIR Light signaling
AT2G32950 ARABIDOPSIS THALIANA CONSTITUTIVE PHOTOMORPHOGENIC 1, ATCOP1, CONSTITUTIVE PHOTOMORPHOGENIC 1, COP1, DEETIOLATED MUTANT 340, DET340, EMB168, EMBRYO DEFECTIVE 168, FUS1, FUSCA 1 TAIR Light signaling
AT2G33835 FES1, FRIGIDA-ESSENTIAL 1 TAIR Vernalization
AT2G34140 TAIR Light signaling
AT2G40080 EARLY FLOWERING 4, ELF4 TAIR Light signaling
AT2G43410 FPA TAIR Autonomous
AT2G46340 SPA1, SUPPRESSOR OF PHYA-105 1 TAIR Light signaling
AT2G46670 TAIR Light signaling
AT2G46790 APRR9, ARABIDOPSIS PSEUDO-RESPONSE REGULATOR 9, PRR9, PSEUDO-RESPONSE REGULATOR 9, TL1, TOC1-LIKE PROTEIN 1 TAIR Light signaling
AT2G46830 ATCCA1, CCA1, CIRCADIAN CLOCK ASSOCIATED 1 TAIR Light signaling
AT2G47700 RED AND FAR-RED INSENSITIVE 2, RFI2 TAIR Light signaling
AT3G02380 ATCOL2, B-BOX DOMAIN PROTEIN 3, BBX3, COL2, CONSTANS-LIKE 2 TAIR Flowering integrator, Light signaling
AT3G04610 FLK, FLOWERING LOCUS KH DOMAIN TAIR Autonomous
AT3G07650 B-BOX DOMAIN PROTEIN 7, BBX7, COL9, CONSTANS-LIKE 9 TAIR Light signaling
AT3G10390 FLD, FLOWERING LOCUS D TAIR Ambient temperature, Autonomous
AT3G12810 CHR13, PHOTOPERIOD-INDEPENDENT EARLY FLOWERING 1, PIE1, SRCAP TAIR Vernalization
AT3G13682 LDL2, LSD1-LIKE2 TAIR Ambient temperature, Autonomous
AT3G15354 SPA1-RELATED 3, SPA3 TAIR Light signaling
AT3G15620 UV REPAIR DEFECTIVE 3, UVR3 TAIR Light signaling
AT3G18990 REDUCED VERNALIZATION RESPONSE 1, REM39, REPRODUCTIVE MERISTEM 39, VRN1 TAIR Vernalization
AT3G20740 FERTILIZATION-INDEPENDENT ENDOSPERM, FERTILIZATION-INDEPENDENT ENDOSPERM 1, FIE, FIE1, FIS3 TAIR Autonomous, Vernalization
AT3G21320 TAIR Light signaling
AT3G24440 VERNALIZATION 5, VIL1, VIN3-LIKE 1, VRN5 TAIR Autonomous, Vernalization
AT3G25730 EDF3, ETHYLENE RESPONSE DNA BINDING FACTOR 3 TAIR Light signaling
AT3G33520 ACTIN-RELATED PROTEIN 6, ARP6, ATARP6, EARLY IN SHORT DAYS 1, ESD1, SUF3, SUPPRESSOR OF FRI 3 TAIR Ambient temperature, Vernalization
AT3G46640 LUX, LUX ARRHYTHMO, PCL1, PHYTOCLOCK 1 TAIR Light signaling
AT3G47500 CDF3, CYCLING DOF FACTOR 3 TAIR Light signaling
AT4G00650 FLA, FLOWERING LOCUS A, FRI, FRIGIDA TAIR Vernalization
AT4G02020 EZA1, SDG10, SET DOMAIN-CONTAINING PROTEIN 10, SWINGER, SWN TAIR Autonomous, Vernalization
AT4G02560 LD, LUMINIDEPENDENS TAIR Autonomous
AT4G08920 ATCRY1, BLU1, BLUE LIGHT UNINHIBITED 1, CRY1, CRYPTOCHROME 1, ELONGATED HYPOCOTYL 4, HY4, OOP2, OUT OF PHASE 2 TAIR Light signaling Grape
AT4G11110 SPA1-RELATED 2, SPA2 TAIR Light signaling
AT4G11880 AGAMOUS-LIKE 14, AGL14 TAIR Vernalization
AT4G16250 PHYD, PHYTOCHROME D TAIR Light signaling
AT4G16280 FCA TAIR Ambient temperature, Autonomous Grape
AT4G16845 REDUCED VERNALIZATION RESPONSE 2, VRN2 TAIR Autonomous, Vernalization
AT4G18130 PHYE, PHYTOCHROME E TAIR Light signaling Grape
AT4G20370 TSF, TWIN SISTER OF FT TAIR Ambient temperature, Flowering integrator
AT4G22950 AGAMOUS-LIKE 19, AGL19, GL19 TAIR Vernalization
AT4G24540 AGAMOUS-LIKE 24, AGL24 TAIR Vernalization
AT4G26000 PEP, PEPPER TAIR Vernalization
AT4G29730 MSI5, NFC5, NUCLEOSOME/CHROMATIN ASSEMBLY FACTOR GROUP C5 TAIR Ambient temperature, Autonomous
AT4G30200 VEL1, VERNALIZATION5/VIN3-LIKE 1, VIL2, VIN3-LIKE 2 TAIR Autonomous, Vernalization
AT4G34530 CIB1, CRYPTOCHROME-INTERACTING BASIC-HELIX-LOOP-HELIX 1 TAIR Light signaling
AT4G35050 MSI3, NFC3, NUCLEOSOME/CHROMATIN ASSEMBLY FACTOR GROUP C 3 TAIR Autonomous, Vernalization
AT4G35900 ATBZIP14, FD, FD-1 TAIR Ambient temperature, Meristem identity
AT5G02810 APRR7, PRR7, PSEUDO-RESPONSE REGULATOR 7 TAIR Light signaling
AT5G03840 TERMINAL FLOWER 1, TFL1 TAIR Ambient temperature, Flowering integrator Grape, Strawberry
AT5G08230 TAIR Vernalization
AT5G08330 CCA1 HIKING EXPEDITION, CHE, TRANSCRIPTION FACTOR TCP21, TCP21 UniProtKB Light signaling
AT5G10140 AGAMOUS-LIKE 25, AGL25, FLC, FLF, FLOWERING LOCUS C, FLOWERING LOCUS F TAIR Ambient temperature, Flowering integrator, Vernalization Grape
AT5G13480 FY TAIR Autonomous
AT5G15840 B-BOX DOMAIN PROTEIN 1, BBX1, CO, CONSTANS, FG TAIR Flowering integrator, Light signaling
AT5G15850 ATCOL1, B-BOX DOMAIN PROTEIN 2, BBX2, COL1, CONSTANS-LIKE 1 TAIR Flowering integrator, Light signaling
AT5G23150 ENHANCER OF AG-4 2, HUA2 TAIR Vernalization
AT5G23280 TAIR Light signaling
AT5G24470 APRR5, PRR5, PSEUDO-RESPONSE REGULATOR 5 TAIR Light signaling
AT5G24930 ATCOL4, B-BOX DOMAIN PROTEIN 5, BBX5, COL4, CONSTANS-LIKE 4 TAIR Light signaling
AT5G35840 PHYC, PHYTOCHROME C TAIR Light signaling Grape
AT5G37055 ATSWC6, SEF, SERRATED LEAVES AND EARLY FLOWERING TAIR Vernalization
AT5G39660 CDF2, CYCLING DOF FACTOR 2 TAIR Light signaling
AT5G42790 ARS5, ARSENIC TOLERANCE 5, ATPSM30, PAF1, PROTEASOME ALPHA SUBUNIT F1 TAIR Vernalization
AT5G48250 B-BOX DOMAIN PROTEIN 8, BBX8 TAIR Light signaling
AT5G57360 ADAGIO 1, ADO1, FKF1-LIKE PROTEIN 2, FKL2, LKP1, LOV KELCH PROTEIN 1, ZEITLUPE, ZTL TAIR Light signaling
AT5G57380 VERNALIZATION INSENSITIVE 3, VIN3 TAIR Autonomous, Vernalization
AT5G57660 ATCOL5, B-BOX DOMAIN PROTEIN 6, BBX6, COL5, CONSTANS-LIKE 5 TAIR Light signaling
AT5G58230 ARABIDOPSIS MULTICOPY SUPRESSOR OF IRA1, ATMSI1, MATERNAL EFFECT EMBRYO ARREST 70, MEE70, MSI1, MULTICOPY SUPRESSOR OF IRA1 TAIR Autonomous, Vernalization
AT5G59570 BOA, BROTHER OF LUX ARRHYTHMO TAIR Light signaling
AT5G60100 APRR3, PRR3, PSEUDO-RESPONSE REGULATOR 3 TAIR Light signaling
AT5G61380 APRR1, ATTOC1, PRR1, PSEUDO-RESPONSE REGULATOR 1, TIMING OF CAB EXPRESSION 1, TOC1 TAIR Light signaling
AT5G62430 CDF1, CYCLING DOF FACTOR 1 TAIR Light signaling
AT5G65050 AGAMOUS-LIKE 31, AGL31, MADS AFFECTING FLOWERING 2, MAF2 TAIR Ambient temperature, Flowering integrator, Vernalization
AT5G65060 AGAMOUS-LIKE 70, AGL70, FCL3, MADS AFFECTING FLOWERING 3, MAF3 TAIR Ambient temperature, Flowering integrator, Vernalization
AT5G65070 AGAMOUS-LIKE 69, AGL69, FCL4, MADS AFFECTING FLOWERING 4, MAF4 TAIR Ambient temperature, Flowering integrator, Vernalization
AT5G65080 AGAMOUS-LIKE 68, AGL68, MADS AFFECTING FLOWERING 5, MAF5 TAIR Ambient temperature, Flowering integrator, Vernalization