Presentation on theme: "Effects of Climate Change on Mechanisms That Initiate Flowering"— Presentation transcript:
1Effects of Climate Change on Mechanisms That Initiate Flowering Donald GeigerUniversity of DaytonMarianist Environmental Education Center
2Basis For Study of Flowering Phenology Exposure to warm temperatures over time is the main driver for springtime plant development, particularly for perennial plants in temperate zones.Plants require a certain amount of exposure to warm temperatures before leaf out or flowering can occur.The required heat sum, a quantified requirement that controls development, is a genetically controlled adaptive trait.One of the most obvious ways ecosystems are affected by global climate change is through alteration of organisms’ developmental timing, their phenology.Onset of flowering is an important element of phenology.
3Climate Change and Flowering Phenology The effect of climate change on the phenology of flower initiation was reported in the following publication: McEwan RW, RJ Brecha, DR Geiger, GP John 2011 Flowering phenology change and climate warming in southwestern Ohio. Plant Ecology 212,(1)The above study, based on observations of first flowering dates for an array of species from 1976 to 2003, provided evidence that climate change is affecting plant development.
4Statisticaly not earlier. Effect of Climate Warming on Flowering PhenologyDevelopmentDevelopmentQuantitative L-DStatisticaly not earlier.DevelopmentWoody OrnamentalQuantitative L-DDevelopmentWoody FruitQuantitative L-DDevelopmentDevelopmentQualitative L-DSpring EphSpring EphIn springIn summerChange in first flowering date over a 28–year (1976 to 2003) observation period in southwestern Ohio, USA. Change in flowering date represents the slope of the linear regression between first flowering date and year. Darkened symbols represent slopes that were different from zero (P<0.05). Horizontal dashed line represents no change.McEwan et al. Plant Ecology (2011) 2112:55-61
5Basis For Flower Phenology Studies This study revealed a trend of progressively earlier flowering time for a number of species.The two earliest flowering species showed the largest advance in flowering time; advance decreased with later flowering date.Two species that bloomed in mid summer also had advanced flowering dates.Flowering is known to be initiated by a number of different physiological mechanisms.Hypothesis: The two clusters of advanced bloom dates may result from climate change acting on two different mechanisms that regulate initiation of flowering.
7Background: A Survey of Physiological Mechanisms of Flowering Phenology In their discussion the authors of the flowering phenology paper “suggest that future work is needed that bridges the gap between pattern (plant phenology response) and process (plant development physiology) in relation to particular climatic cues”.The study that follows is a survey of process… mechanisms that initiate flowering and investigates possible ways in which climate change may advance initiation mediated by these mechanisms… pattern.
9Shoot Apex Stages for Flowering AdultReproductivestateFloral evocation in the adult reproductive phase involves three states on the way to flowering:emphasis on developmental statesCompetent- Shoot apex becomes competent to respond appropriately to a given developmental signal. E.g.- able to respond to photoperiodDetermined- able to follow some developmental program even if shoot is removed from its normal position and nurtured by grafting or rooting E.g.- able to respond to plant growth regulator.Expressed- apical meristem undergoes physiological steps that lead to morphogenesis to become a flower. (Process on slide 10)
10Signal to apex in determined state shoot apexbractetalVegetative shoot apexa.Signal to apex in determined statec.epalstamen primordiashoot pexb.d.Floral Induction and Development of the Flower10
11Survey of Factors Involved In Altering Flowering Time
12Biochemical Signaling Is Able To Initiate Flowering Mikhail Chailakhyan in 1937 demonstrated that floral induction can be transmitted through a graft indicating that a signal substance is being sent.The hypothetical substance was named florigen.In 2005 an RNA was mistakenly identified as florigen In 2007 the paper was retracted; In 2008 florigen was found to be a protein a translocated floral initiation signal, florigen.Florigen proteins FT , Hd3a and PIF4 have been identified as biochemical signals known to be translocated to the apical meristem and to initiate flower formation.12
13Biochemical Signaling Involved in Flowering Mitigating factors:Flowering isexpressed byaction of the florigen signal in response to photoperiod or warming stimulus.Multiple development pathways for floral evocation in Arabidopsis. Part 2
14Some Factors That Initiate Flowering: Candidates for Climate Responses Temperature- warming after chillingTemperature- vernalization = period of chillingPhotoperiod- day length; dark period lengthBiennial life cycle- vernalization followed by lengthening daysCarbohydrate pathway- sucrosePlant growth regulators- gibberellin, ethylene pathway-Autonomous initiation- age, size, leaf number.Significance:- Existence of multiple pathways provides flexibility to produce seeds under a variety of conditions.Which of these mechanisms are affected by climate change?
15Photoperiodism: Monitoring Day Length Day long enoughNight long enough“day”too long“night”too shortLong-Day plantsShort-Day plantsPhotoperiodic regulation of flowering showing effects on short-day plants and long-day plants. Bar graphs show the effects of the duration of dark periods on flowering; data show the critical variable is length of the dark period.
16Biochemical Signaling Involved in Flowering [pp 660 – 667]Long DayShort DayDetermined bywarming signalFlorigen gene expression regulated by warm temperature . PIF4 protein, a florigenDetermined byphotoperiod signalFT protein, the “florigen” is transported to the shoot apex to evoke flowering.16
17Case Studies of Flower Types: 1. Warming Activates Florigen Gene
18Warmth Activates Flower Initiation A period of warming over a period of time drives spring flower development, particularly for perennial plants in temperate zones.Plants require a critical amount of exposure to warm temperatures before leaf out or flowering can occur; required heat sum (RHS)RHS for spring development is a genetically controlled, adaptive trait.Enables plants to survive unpredictable early onset of warmth at the start of the growing season that does not persist.Warmth activates PIF4 gene which acts as a florigen; exerts control over the flowering pathway (slide 16).This mechanism is likely to be encountered in a number of species of plants but has not been studied extensively since its recent discovery (Kumar et al. 2012).
19Mechanisms That Can Modulate Flowering Time The thermal control time can be modulated by conditions that affect the rate of accumulation of the required heat sum. RHSDifferent environments may cause plants to require more or less time to accumulate the effective heat sum under the same input rate.Conditions known to modulate RHS mechanism include:snow pack depth photoperiod winter cold treatmentFor example, photoperiod can delay bud break, protecting against responding to the threat of unusually early but intermittent warm temperature. This effect can complicate RHS/flower induction studies..
20Case Studies of Flower Types: Altered Rate of Flower Development 2. Spring Ephemeral Pattern-Altered Rate of Flower Development
21Spring Ephemeral Flowering Cycle Background: Spring Ephemerals Spring ephemerals are species that have an active spring growth period limited to days… adapted to take advantage of a two-month high-light before canopy closure.Examples: fawn lily, Dutchman’s breeches, crocus, bleeding heart, snow drops, tulip, hyacinth, Cardamine spp (red = reported in McEwan et al.)Annual life cycle, Erythronium americanum- yellow fawn lily17 April19 April20 April21 April22 AprilBackground: Spring Ephemerals
22Spring Ephemeral Developmental Life Cycle Diagramic life cycle of Erythronium americanum .The perennial organ is shown in radial section to show bud development5123467Budsummerautumnwinterspring40 to 60 days above groundSoil temperature drives life cycle development that determines flowering time.
23Spring Ephemeral Life Cycle Species that have a 40 to 60 day growth period in spring.Annual life cycle, Erythronium americanum- yelllow fawn lilyDormancy is broken in autumn; bud and root growth continues through the winter at a very slow rate due to the low soil temperatures. (stages 2, 3).As the soil warms shoot expansion is rapid (stages 3, 4), the long underground growth period ends and the above ground phase begins. This process is hastened by the warming due to climate change.Sexual reproduction takes place over a 5- to 6-day period (stages 4,5,6)Senescence of the above ground plant (stage 6) followed by senescence of the root (stage 7) and a stage of apparent dormancy of the perennial organstage 1)… the end of the 40 – 60 day period. Review previous slide.Flowering time was advanced in crocus (Crocus flavus) and bleeding heart (Dicentra specta sp.)the study of phenology affected by climate warming (McEwan et al. 2010).next year
24Case Studies of Flower Types: 3. Flower Bud Dormancy Broken by Spring Chilling; Woody Perennials and Geophytes
25Breaking of Bud Dormancy: Chilling For a wide range of spring flowering perennials a temperature signal typically breaks dormancy of the renewal (winter) buds.Renewal or winter buds form in early summer after leaf development ends and initiate next spring’s flowering.In woody perennials (apple) formation next year’s leaf and flower primordia occurs in this year’s renewal buds in fall and early winter. !!!Flowering is initiated by chilling during spring warming period.Ex. Woody ornamental:- rhododendron; tree fruit species:- apple, pearFlowering time was advanced in apple (Malus domestica) in the study of phenology affected by climate warming (McEwan et al. 2010)In the geophytes renewal buds form on the roots; flowering is initiated by chilling during the spring warming period.Ex. Geophytes:- tulip, peonie
26Breaking of Bud Dormancy: Chilling In woody perennials breaking of bud dormancy in spring is a factor determining the timing of flowering... Example apple, pearBud dormancy in fall is initiated by low temperatures and short photoperiod. Fruit treesIn spring bud dormancy is broken by environmental signals, especially chilling during the spring warming period.Linear accumulation of chilling hours occurs below 7o C. 45o F.Short photoperiod promotes blooming.Phenology of spring woody perennial flower activation is complex and still not well understood.Remember: flowers formed in late fall, early winter in woody perennials. It is bud break that determined flower activation.
274. Flowering in Qualitative, Obligate Long-Day Plants Case Studies of Flower Types:4. Flowering in Qualitative, Obligate Long-Day Plants
28Qualitative Obligate Long-Day Plants Annual Phlox (Phlox drummondii) is a qualitative long-day plant.qualitative L-D plant:- flowers ONLY on long days… obligate; day period must be at least a certain critical length to initiate floweringquantitative L-D plant:- flowering ACCELERATED by long d. ; facultative:- long days are not required for flowering.Under long-day conditions leaf processes mediated by phytochrome produce florigen that initiates flower formation by the apical meristem. (Revisit slides 16, 15, and 10)At 11 hrD- 98 days to flowering; 13 hrD 87 d, 15 hrD 75 d, 17 hrD 74 dIn Phlox drummondi flower development did not increase beyond 15hr day.Flowering time was advanced in annual phlox (Phlox drummondi) in the study of phenology affected by climate warming (McEwan et al. 2010)28
29Case Studies of Flower Types: 5. Flowering in Quantitative Facultative Long-Day Plants
30Effect of Photoperiod and Temperature on Flowering in Quantitative L-D Plants Shortest time to flowering in Ranunculus asiaticus was achieved under 16 hr photoperiod at 16oC .A quantitative (LD accelerates flowering), facultative (LD not required) long day plantBoth increased temperature + longer daylength reduce time to flowering.The decreased time to flowering by lengthening the photoperiod was due to warming advancing development triggered by photoperiod induction of flowering. Ranunculus is a quantitative long day plant. A day of a certain length is not required for flowering to occur.
31Effect of Photoperiod and Temperature on Flowering in Quantitative L-D Plants Control of time to flowering in Viola:Progress to flowering increases with temperature up to 28oC.This pattern promotes earlier flowering as a result of higher springtime temperatures.Progress to flowering also increases with daylength.Not a photoperiod induction of flowering but the requirement pattern for growth of a quantitative long day plant.Response to longer photoperiod is adaptive, lessening the lag imposed by cooler early season temperatures..
32Effect of a Variety of Factors on Timing of Flowering in in Quantitative L-D Plants A quantitative long-day plant but long photoperiod is not used as a cue. Lengthening days promoter stem elongation.Flowers are formed underground during the previous winter or earlier when the plant forms its ramets.In mountainous habitats the first date of bare ground in spring is a good predictor of first flowering date, reminiscent of the winter/spring maturation of the developing flower stalks..Working hypothesis: increasing soil temperature promotes the formation and maturing of flowers initiated with the formation of ramets in the previous fall/winter.
33Case Studies of Flower Types: 6. Flowering in Biennials
34Biennial Sugar Beet Flowering Biennials: First year rosette of sugar beet is unable to form reproductive shoots during the first year of growth.Duplicate genes BvFT1 and BvFT2, homologs of the FT gene mediate regulation of flowering time: BvFT2 promotes while BvFT1 represses.Winter vernalization lowers expression of BvFT1, which renders shoots competent to reproduce.Exposure to lengthening days (quantitative long day) upregulates BvFT2 and maintains BvFT1 in downregulated state.Subsequent exposure to increasing day length (qualitative long day) causes the shoot to become determined.The dependence on LD puts flower induction outside the springtime window of earlier increased temperatures.
35Case Studies of Flower Types: 7. Flower Initiation Part of a Genetically Determined Development Program
36Flowering Initiated At a Specific Development Stage Some species of plants initiate flowering at a specific node (leaf location on the stem).For a certain variety of cotton it occurs just after nodes number 5 or 6.The program development is driven by heat measured in heat units. (next slide)Determination by stage of plant development seems to be a common mechanism.1st FlowerNode 5Node 6Node 4Node 3Node 2Node 1Vegetative BranchMain Stem
37Flowering Initiated At a Specific Development Stage Table 1. The average number of days and heat units required for various growth stages of cotton in the Mid-South.Modified from Oosterhuis, D.M Growth and development of the cotton plant. In: W.N. Miley and D.M. Oosterhuis (eds) Nitrogen Nutrition in Cotton: Practical Issues. Proc. Southern Branch Workshop for Practicing Agronomists. Publ. Amer. Soc. Agron., Madison, WIGrowth StageDaysHeat Units – DD60sPlanting to Emergence4 to 950 to 60Emergence to First Square27 to 38425 to 475Square to Flower20 to 25300 to 350Planting to First Flower60 to 70775 to 850Flower to Open Boll45 to 65850 to 950Planting to Harvest Ready130 to 1602200 to 2600Daytime high °F + Nighttime low °F = DD602
38Flowering Initiated At a Specific Development Stage Species that initiate flowering at a specific stage of development that is promoted by spring warming would advance flowering as a consequence of climate warming.Species that flower in early spring would be most reliably advanced due to climate warming. Those blooming later likely would show more variability.For tomato plants, the first flower is initiated at node 6 to 8 depending on the earliness of the variety. In a commercial variety of tobacco the first flower was reported to occur at or about node 41.In the phenology study Ageratum, Centauria, Dicentra, Hemerocallis, and Hosta flowering appears to be regulated by developmental stage.
39Summary: Possible Mechanisms for Responses of Flowering To Climate Change
40Flowering Phenology and Climate Warming McEwan R, R Brecha, D Geiger, G John 2011 Flowering Phenology Change and Climate Warming in Southwestern Ohio. Plant Ecol 212:Global climate change affects timing of plant development.Thirty years of annual botanical surveys were used to examine impacts of climate change on plant development.Authors of the study found a significant relationship between warming winter and spring temperatures and first flowering time for 60% of the species studied.
41Statisticaly not earlier. Effect of Climate Warming on Flowering PhenologyDevelopmentDevelopmentQuantitative L-DStatisticaly not earlier.DevelopmentWoody OrnamentalQuantitative L-DDevelopmentWoody FruitQuantitative L-DDevelopmentDevelopmentQualitative L-DSpring EphSpring EphIn springIn summerChange in first flowering date over a 28–year (1976 to 2003) observation period in southwestern Ohio, USA. Change in flowering date represents the slope of the linear regression between first flowering date and year. Darkened symbols represent slopes that were different from zero (P<0.05). Horizontal dashed line represents no change.McEwan et al. Plant Ecology (2011) 2112:55-61
42Seven Flower Induction Mechanisms 1. Warming activates florigen gene.Arabidopsis thaliana; newly discovered; not researched2. Spring ephemeral pattern of flower development.Erythronium americanum, Cardamine sp., Galanthus sp., Allium sp., Anemone sp., Crocus sp.3. Flower bud summer dormancy broken in spring by chilling.Woody trees and shrubs: Malus sp. Rhododendron sp.Geophytes: Paeonia, Tulipa4. Long days required in qualitative, obligate long-day plantsPhlox sp.5. Long days accelerate flowering; quantitative L-D plantsRanunculus asiaticus, Viola sp, Delphinium sp.6. Biennial species, chilling period and lengthening daysBeta vulgaris7. Development + warming hastens flowering.Dicentra, Hemerocalis, Hosta
44Did Not AdvanceMar 27: Forsythia sp- Flower bud dormancy broken by chilling.Apr 23: Ajuga reptans- Development ??Apr 27: Geranium maculatum- facult L-D plant; LD + warmingMay 5: Ranunculus sp- DevelopmentMay 13: Centaurea cyanus- DevelopmentAug 8: Ageratum sp- Development
45References [Pg 1] Flowering physiology Taiz L, E Zeiger 2006 Plant Physiology 4th Ed. Sinauer Associates Inc.Zeevaart, JAD Leaf-produced floral signals. Current Opinion in Plant Biology 11:541–547.McDaniel CN, SR Singer and SME Smith Developmental states associated with the floral transition. Developmental Biology 153:59-69.Flowering phenology and climate warmingMcEwan RW, RJ Brecha, DR Geiger, GP John 2011 Flowering phenology change and climate warming in southwestern Ohio. Plant Ecology 212,(1)Conover D, S Pelikan 2010 Earlier flowering in a restored wetland-prairie correlate with warmer temperatures (Ohio). Ecological Restoration. 28(4) :Bertin RI 2008 Plant phenology and distribution in relation to recent climate change. J. Torrey Bot. Soc. 135, 126–146.Lapointe L (2001) How phenology influences physiology in deciduous forest spring ephemerals. Phsiologia Plantarum 113:Bradley NL, AC Leopold, J Ross,. W huffaker 1999 Phenological changes reflect climate change in Wisconsin. Proceedings of the National Academy of Science USA 96:Floral induction mechanism triggered by warmingKumar SV, Lucyshyn D, Jaeger KE, Alos E., Alvev E, Harberd NP, Wigge PA Transcription factor PIF4 controls the thermosensory activation of flowering Nature March
46References [Pg 2] Release of bud dormancy in woody perennials Arora A, Rowland LJ and Tanino K 2003 Induction and release of bud dormancy in woody perennials: a science comes of age. Hort Science 38:Photoperiod and flowering phenologyKarlsson, M 1996 Control of Ranunculus asiaticus flowering by photoperiod and temperature. Hort Science 31:680-68Adams SR, Pearson S and Hadley P 1997 The effects of temperature, photoperiod and light integral on the time to flowering of Pansy cv, Universal Violet. Annals Bot 80;Baloch J, M Munir, M Abid and M Iqbal Effects of different photoperiods on flowering time of qualitative long-day ornamental annuals. Pakistan J Bot.43(3):Modulation of flower induction mechanismsInouye et al Oecologica 130: investigate the roles of snowpack, frost, La Nina, climate change in modulating response.Genetic and physiological bases for phenological responses to climatesWilczek AM,LT Burghardt, AR Cobb, MD Cooper, SM Welch, J Schmidt 2010 Genetic and physiological bases for phenological responses to current and predicted climates. Phil Trqns R Soc B 365 (1555):