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Effects of Climate Change on Mechanisms That Initiate Flowering Donald Geiger University of Dayton Marianist Environmental Education Center.

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Presentation on theme: "Effects of Climate Change on Mechanisms That Initiate Flowering Donald Geiger University of Dayton Marianist Environmental Education Center."— Presentation transcript:

1 Effects of Climate Change on Mechanisms That Initiate Flowering Donald Geiger University of Dayton Marianist Environmental Education Center

2 2 Exposure to warm temperatures over time is the main driver for springtime plant development, particularly for perennial plants in temperate zones 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. 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. 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. 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. Onset of flowering is an important element of phenology. Basis For Study of Flowering Phenology

3 3 Climate Change and Flowering Phenology McEwan RW, RJ Brecha, DR Geiger, GP John 2011 Flowering phenology change and climate warming in southwestern Ohio. Plant Ecology 212,(1) 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.

4 Change 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. 4 Spring Eph Woody Fruit Quantitative L-D McEwan et al. Plant Ecology (2011) 2112:55-61 In spring In summer Effect of Climate Warming on Flowering Phenology Statisticaly not earlier. Development Development Qualitative L-D Development Woody Ornamental Development Quantitative L-D Development Development

5 5 Basis For Flower Phenology Studies progressively earlier flowering time for a number of species This study revealed a trend of progressively earlier flowering time for a number of species. two earliest flowering species showed the largest advance in flowering time The two earliest flowering species showed the largest advance in flowering time; advance decreased with later flowering date. bloomed in mid summer also had advanced flowering dates. Two species that bloomed in mid summer also had advanced flowering dates. different physiological mechanisms. 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. Hypothesis: The two clusters of advanced bloom dates may result from climate change acting on two different mechanisms that regulate initiation of flowering.

6 6 Physiology of Floral Induction:

7 7 Background: 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.

8 Floral Induction Shoot Apex Phase Changes three developmental phases Shoot apical meristems have three developmental phases: 1. Juvenile phase 2. Adult vegetative phase 3. Adult reproductive phase Flower formation

9 9 Shoot Apex Stages for FloweringAdultReproductive state state emphasis on developmental states adult reproductive phase three states Floral evocation in the adult reproductive phase involves three states on the way to flowering: Shoot apex becomes competent to respond appropriately to a given developmental signal. able to respond to photoperiod Competent- Shoot apex becomes competent to respond appropriately to a given developmental signal. E.g.- able to respond to photoperiod able to follow some developmental program even if shoot is removed from its normal position and nurtured by grafting or rooting. able to respond to plant growth regulator. Determined- 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. apical meristem undergoes physiological steps that lead to morphogenesis to become a flower. (Process on slide 10) Expressed- apical meristem undergoes physiological steps that lead to morphogenesis to become a flower. (Process on slide 10)

10 bract shoot apex epal shoot pex etal a. b. c. d. stamen primordia Floral Induction and Development of the Flower Signal to apex in determined state Vegetative shoot apex 10

11 11 Survey of Factors Involved In Altering Flowering Time

12 12 floral induction Mikhail Chailakhyan in 1937 demonstrated that floral induction can be transmitted through a graft indicating that a signal substance is being sent. Mikhail Chailakhyangraft 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. FTHd3aPIF4 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. Biochemical Signaling Is Able To Initiate Flowering

13 13 Biochemical Signaling Involved in Flowering Multiple development pathways for floral evocation in Arabidopsis. Part 2 Flowering is expressed by florigen photoperiodwarming stimulus action of the florigen signal in response to photoperiod or warming stimulus. Mitigating factors:

14 14 warming Temperature- warming after chilling =chilling Temperature- vernalization = period of chilling day lengthdark period length Photoperiod- day length; dark period length vernalization followed by lengthening days Biennial life cycle- vernalization followed by lengthening days sucrose Carbohydrate pathway- sucrose gibberellin, ethylene Plant growth regulators- gibberellin, ethylene pathway- age, size, leaf number Autonomous initiation- age, size, leaf number. Existence of multiple pathways provides flexibility to produce seeds under a variety of conditions Significance:- Existence of multiple pathways provides flexibility to produce seeds under a variety of conditions. Which of these mechanisms are affected by climate change? Some Factors That Initiate Flowering: Candidates for Climate Responses

15 Photoperiodism: Monitoring Day Length 15 effects on short-day plants and long-day plants. data show the critical variable is length of the dark period Photoperiodic 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. Day long enoughNight long enough Short-Day plants Long-Day plants “day” “night” too long too short

16 16 Biochemical Signaling Involved in Flowering [ pp 660 – 667 ] FT protein, the “florigen” is transported to the shoot apex to evoke flowering. Determined by photoperiod signal 16 Determined by warming signal Florigen gene expression regulated by warm temperature. PIF4 protein, a florigen Long Day Short Day

17 17 Case Studies of Flower Types: 1. Warming Activates Florigen Gene

18 18 Warmth Activates Flower Initiation A period of warming over a period of time A period of warming over a period of time drives spring flower development, particularly for perennial plants in temperate zones. require a critical amount of exposure to warm temperatures required heat sum (RHS) Plants require a critical amount of exposure to warm temperatures before leaf out or flowering can occur; required heat sum (RHS) RHSgenetically controlled, adaptive trait. RHS for spring development is a genetically controlled, adaptive trait. E Enables plants to survive unpredictable early onset of warmth at the start of the growing season that does not persist. slide 16 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).

19 19 Mechanisms That Can Modulate Flowering Time The thermal control time modulated conditions rate of accumulation required heat sumRHS The thermal control time can be modulated by conditions that affect the rate of accumulation of the required heat sum. RHS effective heat sum under the same input rate. Different environments may cause plants to require more or less time to accumulate the effective heat sum under the same input rate. Cmodulate RHS mechanism Conditions known to modulate RHS mechanism include: snow pack depth photoperiod winter cold treatment snow pack depth photoperiod winter cold treatment photoperiod For 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..

20 20 Case Studies of Flower Types: 2. Spring Ephemeral Pattern- Altered Rate of Flower Development

21 21 Background: Spring Ephemerals adapted to take advantage of a two-month high-light before canopy closure 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. crocus, bleeding heart, snow drops red = reported in McEwan et al. 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 lily 17 April 20 April 19 April 21 April 22 April Spring Ephemeral Flowering Cycle

22 40 to 60 days above ground Spring Ephemeral Developmental Life Cycle Diagramic life cycle of Erythronium americanum. The perennial organ is shown in radial section to show bud development summer autumn winter spring Bud Soil temperature drives life cycle development that determines flowering time. 22

23 23 Spring Ephemeral Life Cycle Species that have a 40 to 60 day growth period in spring. Annual life cycle, Erythronium americanum- yelllow fawn lily Dormancy is broken in autumn; (stages 2, 3) Dormancy 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). shoot expansion is rapid (stages 3, 4) above ground phase begins 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 (stages 4,5,6) Sexual reproduction takes place over a 5- to 6-day period (stages 4,5,6) Senescence of the above ground plant (stage 6) (stage 7) apparent dormancy of the perennial organ 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 organ stage 1)… the end of the 40 – 60 day period.Review previous slide. stage 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

24 24 Case Studies of Flower Types: 3. Flower Bud Dormancy Broken by Spring Chilling; Woody Perennials and Geophytes

25 25 Breaking of Bud Dormancy: Chilling For a wide range of spring flowering perennials a temperature signal typically breaks dormancy of the renewal (winter) buds. 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. 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. !!! 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. Flowering is initiated by chilling during spring warming period. Ex. Woody ornamental:- rhododendron; tree fruit species:- apple, pear Flowering 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. In the geophytes renewal buds form on the roots; flowering is initiated by chilling during the spring warming period. Geophytes:- Ex. Geophytes:- tulip, peonie

26 26 breaking of bud dormancy in spring In woody perennials breaking of bud dormancy in spring is a factor determining the timing of flowering... Example apple, pear low temperatures and short photoperiodFruit trees Bud dormancy in fall is initiated by low temperatures and short photoperiod. Fruit trees especially chilling during the spring warming period. In spring bud dormancy is broken by environmental signals, especially chilling during the spring warming period. Linear accumulation of chilling hours occurs below 7 o C. 45 o F. Linear accumulation of chilling hours occurs below 7 o C. 45 o 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. Remember: flowers formed in late fall, early winter in woody perennials. It is bud break that determined flower activation. Breaking of Bud Dormancy: Chilling

27 27 Case Studies of Flower Types: 4. Flowering in Qualitative, Obligate Long-Day Plants

28 28 Qualitative Obligate Long-Day Plants 28 Phloxqualitative Annual Phlox (Phlox drummondii) is a qualitative long-day plant. qualitativeONLY on long daysobligate; at least a certain critical length qualitative L-D plant:- flowers ONLY on long days… obligate; day period must be at least a certain critical length to initiate flowering quantitativeACCELERATED by long d. ; facultative:- long days are not required for flowering. quantitative L-D plant:- flowering ACCELERATED by long d. ; facultative:- long days are not required for flowering. slides 16, 15, 10) 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 d Phlox drummondi In 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)

29 29 Case Studies of Flower Types: 5. Flowering in Quantitative Facultative Long-Day Plants

30 30 16 hr photoperiod at 16 o C. Shortest time to flowering in Ranunculus asiaticus was achieved under 16 hr photoperiod at 16 o C. quantitative (LD accelerates flowering), facultative (LD not required) long day plant A quantitative (LD accelerates flowering), facultative (LD not required) long day plant increased temperature + daylength Both 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. 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. Effect of Photoperiod and Temperature on Flowering in Quantitative L-D Plants

31 31 Viola Control of time to flowering in Viola: Progress to flowering increases with temperature up to 28 o C. This pattern promotes earlier flowering as a result of higher springtime temperatures. This pattern promotes earlier flowering as a result of higher springtime temperatures. Progress to flowering also increases with daylength. Progress to flowering also increases with daylength. the requirement pattern for growth of a quantitative long day plant. 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. Response to longer photoperiod is adaptive, lessening the lag imposed by cooler early season temperatures.. Effect of Photoperiod and Temperature on Flowering in Quantitative L-D Plants

32 32 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 ramets 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.. increasing soil temperature promotes the formation and maturing of flowers initiated with the formation of ramets in the previous fall/winter. Working hypothesis: increasing soil temperature promotes the formation and maturing of flowers initiated with the formation of ramets in the previous fall/winter. Effect of a Variety of Factors on Timing of Flowering in in Quantitative L-D Plants

33 33 Case Studies of Flower Types: 6. Flowering in Biennials

34 34 unable to form reproductive shoots Biennials: First year rosette of sugar beet is unable to form reproductive shoots during the first year of growth. BvFT1BvFT2 BvFT2promotesBvFT1represses Duplicate genes BvFT1 and BvFT2, homologs of the FT gene mediate regulation of flowering time: BvFT2 promotes while BvFT1 represses. vernalizationBvFT1 competent Winter vernalization lowers expression of BvFT1, which renders shoots competent to reproduce. quantitative long day) BvFT2 Exposure to lengthening days (quantitative long day) upregulates BvFT2 and maintains BvFT1 in downregulated state. qualitative long day) determined S ubsequent 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. Biennial Sugar Beet Flowering

35 35 Case Studies of Flower Types: 7. Flower Initiation Part of a Genetically Determined Development Program

36 36 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. Flowering Initiated At a Specific Development Stage Node 4 Node 3 Node 2 Node 1 Node 5 Node 6 1 st Flower Main Stem Vegetative Branch

37 37 Flowering Initiated At a Specific Development Stage Growth StageDaysHeat Units – DD60s Planting to Emergence4 to 950 to 60 Emergence to First Square27 to to 475 Square to Flower20 to to 350 Planting to First Flower60 to to 850 Flower to Open Boll45 to to 950 Planting to Harvest Ready130 to to 2600 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, WI Daytime high °F + Nighttime low °F -60 = DD60 2

38 38 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. Flowering Initiated At a Specific Development Stage

39 39 Summary: Possible Mechanisms for Responses of Flowering To Climate Change

40 40 Flowering Phenology and Climate Warming McEwan R, R Brecha, D Geiger, G John 2011 Plant Ecol 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.

41 Change 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. 41 Spring Eph Woody Fruit Quantitative L-D McEwan et al. Plant Ecology (2011) 2112:55-61 In spring In summer Effect of Climate Warming on Flowering Phenology Statisticaly not earlier. Development Development Qualitative L-D Development Woody Ornamental Development Quantitative L-D Development Development

42 42 1. Warming activates florigen gene. Arabidopsis thaliana; newly discovered; not researched 2. 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, Tulipa 4Long days required qualitative, obligate long-day plants 4. Long days required in qualitative, obligate long-day plants Phlox sp. 5. Long days accelerate flowering; quantitative L-D plants, Ranunculus asiaticus, Viola sp, Delphinium sp. 6. Biennial species, chilling period and lengthening days Beta vulgaris 7. Development + warming hastens flowering. Dicentra, Hemerocalis, Hosta Seven Flower Induction Mechanisms

43 43 Advanced Spring ephemeral Mar 2: Galanthus sp.- Spring ephemeral Spring ephemeral Mar 6: Crocus flavus- Spring ephemeral Quantitative L-D plant; LD? + warming Apr 18: Viola pubescens- Quantitative L-D plant; LD? + warming Woody fruit, bud dormancy broken by chilling Apr 19: Malus sp.- Woody fruit, bud dormancy broken by chilling Quant. L-D plant; LD? + warming Apr 25: Delphinium consolida- Quant. L-D plant; LD? + warming Quant L-D plant; LD? + warming Apr 27: Dicentra spectabilis- Quant L-D plant; LD? + warming Development;Dev + warm May 19: Hemerocalus lilioasphodelus- Development; Dev + warm Qualitative, long-day plant; LD + warming Jul 14: Phlox sp- Qualitative, long-day plant; LD + warming Development;Dev + warm Aug 11: Hosta plantaginea- Development; Dev + warm

44 44 Did Not Advance Flower bud dormancy broken by chilling. Mar 27: Forsythia sp- Flower bud dormancy broken by chilling. Development ?? Apr 23: Ajuga reptans- Development ?? facult L-D plant; LD + warming Apr 27: Geranium maculatum- facult L-D plant; LD + warming Development May 5: Ranunculus sp- Development Development May 13: Centaurea cyanus- Development Development Aug 8: Ageratum sp- Development

45 45  Flowering physiology  Taiz L, E Zeiger 2006 Plant Physiology 4 th 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:  Flowering phenology and climate warming  McEwan 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 warming  Kumar SV, Lucyshyn D, Jaeger KE, Alos E., Alvev E, Harberd NP, Wigge PA Transcription factor PIF4 controls the thermosensory activation of flowering Nature March References [Pg 1]

46 46  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 phenology  Karlsson, M 1996 Control of Ranunculus asiaticus flowering by photoperiod and temperature. Hort Science 31:  Adams 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 mechanisms  Inouye 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 climates  Wilczek 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): References [Pg 2]


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