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The Reproductive Phenology of Citrus. II: Citrus Floral Ontogeny

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The Reproductive Phenology of Citrus. II: Citrus Floral Ontogeny The reproductive phenology of Citrus. II: Citrus floral ontogeny JAKKIE (OPJ) STANDER Citrus Research International, Department of Horticultural Science, University of Stellenbosch, Private Bag X1, 7602, Matieland. E-mail: [email protected] OPSOMMING: • Hoё vlakke van interne ouksiene (IAA) three articles on the physiology of citrus • Meerderheid sitrus-spesies blom gedurende afkomstig van vrugte, inhibeer die vorming flowering, and will focus on the important die lente. Die verskynsel word voorafgegaan van nuwe blom dra-posisies gedurende die chronological events pertaining to the devel- deur ʼn blom-ontwikkelingsproses gedurende somer. opment of a citrus flower, as influenced by die herfs en winter (Mei-Julie). • Vroeё verwydering van vrugte (uitdunning endogenous factors, environmental condi- • Blom-induksie (BI) is die eerste en bepalen- in “aan”-jaar) verminder die inhiberende ef- tions and cultural practices. de stap in die blom-ontwikkelingsproses. fek van ouksiene op ontwikkeling van nuwe • Water stres inisieёr BI in warm, somer- lote (apikale dominansie) en stimuleer die Inleiding reёnval streke en genoegsame lae tempera- vorming van nuwe dra-posisies. Sitrus is ʼn immergroen boom wat ʼn kom- ture (15-20°C) in koeler, winter-reёnval • Lae koolhidraat-vlakke gedurende periode plekse boomstruktuur onderhou deur een streke. van blom-ontwikkeling beperk die potensi- tot drie jaarlikse vegetatiewe groei-fases. Na • Die belangrike FT-geen, is onlangs in aal vir blom-knop ontwikkeling en kan lei tot induksie van nuutgevormde vegetatiewe lote blom-plante geidentifiseer en sy uitdrukking swak blom-ontwikkeling. deur genoegsame geakkumuleerde lae tem- is direk gekoppel aan tyd van blom, sowel as • Verwydering van vrugte gedurende ʼn aan- perature en/of water stress gedurende die blom-intensiteit. jaar deur chemiese- of handuitdunning is ʼn herfs en winter, ontwikkel blomme in die • Water stres en/of lae temperature lei tot die suskesvolle manier om ʼn balans tussen bel- lente vanuit knoppe van lote wat gedurende uitdrukking van die sitrus FT-geen (CiFT) in angrike interne hormone soos GA en IAA te die vorige 12-maand seisoen gevorm het. blare, lote en knoppe tydens BI. Die CiFT- handhaaf, asook om die opbou van koolhi- Hierdie artikel is die tweede in ʼn reeks mRNA word transleer na CiFT-proteien, drate tot ʼn genoegsame vlak te bewerkstellig. van drie artikels wat handel oor die fisiolo- wat daarna in die phloeёm van die loot na • Bemesting en besproeingspraktyke ge- gie van blom-ontwikkeling in sitrus en die ontvanklike knoppe vervoer word. Genoeg- durende die somer en herfs moet so aangepas belangrike stappe van blom-ontwikkeling, same akkumulasie van die proteien in die word om nuwe vegetatiewe groei te stimuleer soos beinvloed deur interne plant-kondisies, knop, onder omgewingskondisies gunstig vir en sodoende nuwe blom dra-posisies te be- omgewingsfaktore en produksie-praktyke. groei, lei tot blom-ontwikkeling gedurende werkstellig. die lente. Flower induction • Natuurlike interne gibbereliene afkomstig Introduction In subtropical climates, citrus flowering (with van sade en die skil van vrugte onderdruk Citrus trees are perennial evergreens, which the exception of lemon and lime) occurs dur- die uitdrukking van die CiFT-geen en die sustain a complex tree structure with one ing spring and is preceded by an intricate, produksie van die CiFT-proteien. Dieselfde to three distinct annual vegetative growth synchronized flower development process respons word verkry van blaartoedienings flushes. After a sufficient induction period, during autumn and winter (April-July) (Kra- van sintetiese gibbereliene (GA) gedurende with the onset of growth-promoting condi- jewski and Rabe, 1995). Flower induction is herfs en winter. tions during spring (increase in temperatures the first and essential step in this process and • In kultivars wat laat in die seisoen ge-oes and sufficient water and mineral nutrient precedes the first microscopic signs of flower word, kan die aanwesigheid van ʼn uiter- supply), flowers develop from buds on veg- initiation by 3 to 4 weeks (Monselise and matige hoё aantal GA-ryke vrugte (“aan”- etative shoots that originated from vegetative Halevy, 1964). Citrus is day-length neutral jaar) die ekspressie van die CiFT-geen on- shoots in the different growth flushes of the (Davenport, 1990; Moss, 1969), with the pre- derdruk. Dit kan lei tot die ontwikkeling van previous 12-month season. dominant promoting stimulus for flower in- min of geen blomme. This article is the second in a series of duction in warm, summer-rainfall areas be- TECHNOLOGY | CRI 65 AUGUST | SEPTEMBER 2015 currently provides a viable model of flower induction in both annual/biennial and per- ennial flowering plants. The accepted model states that FT-gene expression in plant leaves in response to an environmental cue results in FT-protein synthesis and transport via the phloem to a susceptible meristem. Bud tran- Figure 1. Citrus trees develop new potential flowering shoots during vegetative growth flushes sition from a vegetative to reproductive state in the spring, summer and autumn (1). Sufficient hours of cool temperature (15-20°C) and/or only initiates under growth-promoting envi- a prolonged period of water stress, lead to the expression of the citrus FT-gene (CiFT) in leaves ronmental conditions such as warm temper- and buds on the newly developed shoots (2A). The CiFT-mRNA is translated to the CiFT-protein atures and increased water supply, once the and transported via the phloem to susceptible buds (2B). Under growth-promoting conditions, accumulation of FT-proteins in the bud have a threshold level of CiFT-protein will result in development of flower parts within the bud and reached a certain species-specific threshold sprouting of flowering shoots. level (Nishikawa et al., 2007). In citrus, a sufficient period of cool tem- perature (15-20°C) and/or a prolonged pe- riod of water stress, are the main inductive stimuli that lead to expression of the citrus FT-gene (CiFT) in leaves, stems and buds (Nishikawa et al., 2007) (Fig. 1). After suffi- cient induction, the CiFT-mRNA is translat- ed to the CiFT-protein, which is transported via the phloem to susceptible buds, and re- sults in reproductive development under growth-promoting conditions during spring (Fig. 1). Loss of function in the CiFT-gene delays flowering, whereas over-expression Figure 2. Fruit rinds and developing seeds are major sources of endogenous hormones. Auxins leads to very early and/or intense flowering (IAA) transmitted within the shoot during summer, reduce the sprouting of lateral buds and (Nishikawa et al., 2010). development of new potential flowering shoots (1). During flower induction in the winter, gibberellins (GAs) reduce the expression of the key flower promoting gene, the citrus flowering Flower initiation and locus T (CiFT) in the leaves and buds (2A+B) and results in a lack of flowers during spring (3). differentiation Flower initiation in citrus involves the tran- ing that of water stress (Moss, 1969) and low understanding flower induction has recently sition of bud meristematic tissue from a veg- temperatures in cooler, winter-rainfall areas come about with the successful identifica- etative to reproductive state in response to (Valiente and Albrigo, 2004). Both flower- tion of a key flowering gene, the FLOWER- levels of sufficiently accumulated CiFT-pro- promoting stimuli result in the cessation ING LOCUS T (“FT-gene”) in the model teins in the bud (Davenport, 1990). Flower of root growth, which strongly resembles a plant, Arabidopsis, and with the expression initiation is the process within the bud, dur- common mechanism of flower induction in- of its homologues and the direct relationship ing which the flower parts start to develop itiation. For many years, the hypothesis sur- to flowering also being confirmed in peren- at a molecular level into a state significantly rounding this common mechanism was that nial fruit tree species such as citrus (Nishi- distinguishable from vegetative, non-flower- of an obligatory/required tree “rest” period, kawa et al., 2007). The intensity of FT-gene ing buds (Lord and Eckard, 1985). The dif- similar to that of winter dormancy in pome expression in reaction to species-specific en- ferentiation of a bud occurs with the onset of and stone fruit species. vironmental stimuli strongly correlates with growth-promoting conditions during spring However, a significant breakthrough in time of flowering and flower intensity, and and results in bud transformation into either TEGNOLOGIE | CRI 66 AUGUSTUS | SEPTEMBER 2015 a vegetative or reproductive state. Non-dif- cultivar differences in response to climate In addition, summer fruit removal (hand ferentiating buds remain dormant as a result and water supply (Davenport, 1990), and thinning) results in a significant increase in of sprouting inhibition incurred by presence time of harvest (early vs. late) (Verreynne and the size of remaining fruit (Stander, 2014) of fruit (Verreynne and Lovatt, 2009), insuf- Lovatt, 2009) all point to the presence of an and results in the decrease in an important ficient growth-promoting conditions (Moss, endogenous, self-regulating system that per- group of endogenous hormones, gibberellins 1969) or age of the bud/shoot (Schneider, tains to the intensity of flowering response. (GA), which directly influence flower-bud 1968). Concomitantly, the majority of research development at the molecular level. Flowering buds differentiate first - ap- findings
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