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Floral Ontogeny of Pelargonium ´Domesticum

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Floral Ontogeny of Pelargonium ´Domesticum J. AMER. SOC. HORT. SCI. 125(1):36–40. 2000. Floral Ontogeny of Pelargonium ×domesticum Marietta Loehrlein1 and Richard Craig2 Department of Horticulture, The Pennsylvania State University, University Park, PA 16802 ADDITIONAL INDEX WORDS. regal pelargonium, geranium ABSTRACT. Floral ontogeny of two cultivars of Pelargonium ×domesticum L.H. Bailey, (regal pelargonium) ‘Duchess’ and ‘Jennifer’, was examined. Plants of both cultivars were grown together in a growth chamber at 15.5 °C with a photosynthetic photon flux of 10 mol·m–2·d–1. Meristems were examined at 5-day intervals over an experimental period of 170 days. The initial vegetative meristem was convex with leaf primordia initiated on either side in an alternate pattern. Early floral initiation was characterized by formation of two clefts on either side of the meristem. Between the clefts new meristems developed. Proliferation of meristems continued until numerous meristems were organized in a cluster arrangement at the apex of the shoot. New meristems lacked leaf primordia and would develop into flowers. Floral organ primordia on a floral meristem were initiated in a succession of four whorls: sepals, petals, androecia, and gynoecium. Pelargonium ×domesticum (regal pelargonium) is a day- nium) (this species is also known in the horticultural literature as neutral plant (Post, 1949). Previous studies have demonstrated Pelargonium zonale) are available (Sattler, 1973; Wetzstein and that development of visible flower buds in regal pelargonium is Armitage, 1983). However, there is no published description of controlled by both temperature and irradiance (Hackett and the floral ontogeny of Pelargonium ×domesticum. Therefore, the Kister, 1974; Nilsen, 1975; Post, 1949). However, development objective of this study was to examine and describe the floral of visible flower buds is a relatively late stage in the transition of ontogeny of regal pelargonium meristems under a controlled set a vegetative meristem to an inflorescence and the subsequent of environmental conditions. development of floral organs. Preceding this development, mo- lecular changes occur in the plant which signal changes in the Materials and Methods meristem (induction) (Bernier, 1986; Bernier et al., 1993). Mo- lecular and cellular changes occur in the meristem in response to PLANT MATERIALS AND CULTURE. Six-week-old cuttings of two the inductive signal (evocation). As cells respond to molecular cultivars of P. ×domesticum, ‘Duchess’ and ‘Jennifer’, were ob- signals, cellular reorganization occurs in the meristem. The result tained from Oglevee, Ltd. (Connellsville, Pa.), on 19 Dec. 1996. The is a change in morphogenesis of the meristem and commence- rooted cuttings were planted into Rubber Dirt plugs (Oglevee, Ltd.), ment of floral initiation (Huala and Sussex, 1993; Sattler, 1973; in a styrofoam Pod Pack (Oglevee Ltd.). Rubber Dirt is composed Steeves and Sussex, 1994). The sequence of events in a plant as of peat moss, water, and a hydrophilic polymer. Each of four pod the meristem changes from indeterminate (vegetative) to deter- packs contained 26 plants, for a total of 104 cuttings of each cultivar. minate (floral) is vegetative → induction → evocation → initia- The two cultivars had been chosen based on several criteria: 1) they tion → organogenesis → floral organ development. are modern commercial cultivars; 2) they were developed under Since both temperature and irradiance affect development of different environmental conditions (‘Duchess’ was developed in visible flower buds in regal geranium (Hackett and Kister, 1974; England by Ernest Walters and ‘Jennifer’ was developed in Califor- Nilsen, 1975; Post, 1949), it is possible that different environ- nia by David Lemon of Oglevee, Ltd.); 3) they were developed from mental factors are important at different stages of floral ontogeny. different germplasm sources; and 4) they have different total Therefore, it is desirable to examine each stage of floral ontogeny cumulative irradiance requirements for floral initiation: ‘Duchess’ under a controlled set of environmental factors. In the future it cuttings require ≈200 mol and ‘Jennifer’ cuttings require ≈300 mol will be possible to alter environmental factors and examine the (Loehrlein, 1997). effects, if any, on floral ontogeny compared to the set of environ- ENVIRONMENTAL CONDITIONS. Upon arrival, rooted cuttings mental factors used in this study. were transferred to a growth chamber (model GC15; Environ- Descriptions of the floral ontogeny of Pelargonium inquinans mental Growth Chamber, Chagrin Falls, Ohio) with temperature L. (Payer, 1857) and P. ×hortorum L.H. Bailey (bedding gera- and humidity set at 15.5 °C and 30%, respectively. A relatively low humidity was desired to prevent infection by fungal patho- Received for publication 22 Dec. 1998. Accepted for publication 11 Oct. 1999. gens. Irradiance was provided at a photosynthetic photon flux This work was supported in part by funding from the Fred C. Gloeckner (PPF) of 175 µmol·m–2·s–1 for a 16-h photoperiod, with a daily Foundation and the Pennsylvania State University Intercollegiate Graduate light integral of 10 mol·m–2. Irradiance was provided by 16 cool- Program in Genetics. We would especially like to thank Oglevee, Ltd., Connellsville, Pa., for providing plant material. We would also like to express white fluorescent (F72T12CWHO; Kloninklijke Philips Elec- gratitude to Angelica Zhigelei and Ken Myers for technical support. From a tronics, N.V., Eindhoven, The Netherlands), high output 160-W dissertation submitted by the senior author in partial fulfillment of the require- lamps. Ten percent of the input wattage was provided by 60-W ments for the PhD degree in genetics. Scanning electron microscopy was incandescent bulbs (Philips 60 W, 130 v). Irradiance was mea- performed at the Pennsylvania State University Scanning Electron Microscope Facility for Life Sciences in the Biotechnology Institute. The cost of publishing sured using a light meter fitted with a quantum sensor (LI-185A; this paper was defrayed in part by the payment of page charges. Under postal LI-COR, Lincoln, Neb.) to measure PPF. Readings were made at regulations, this paper therefore must be hereby marked advertisement solely to six locations within the chamber, and an overall average was then indicate this fact. calculated. At the center of the chamber irradiance was 200 1 Current address: Assistant professor of horticulture, Department of Agriculture, µmol·m–2·s–1, while near the outer edges, irradiance was 150 Western Illinois University, Macomb, IL 61455. µ –2 –1 2J. Franklin Styer professor of horticultural botany, Department of Horticulture, mol·m ·s PPF. To provide all experimental units with uni- Pennsylvania State University, University Park, PA 16802. form exposure to irradiance, pod packs were rotated daily. 36 J. AMER. SOC. HORT. SCI. 125(1):36–40. 2000. Plants were checked daily and fertigated when media began to Results dry with 1.33 g·L-1 Excel 15–2.2–12.5 Cal-Mag water-soluble fertilizer (Scotts-Sierra Horticultural Products, Marysville, Ohio). FLORAL MORPHOLOGY. Florets of Pelargonium ×domesticum Plants were irrigated once a week with tap water to reduce salt were five-merous, perfect and complete. They were borne in a accumulation. compound umbel. Petiolate florets formed secondary umbels, EXPERIMENTAL DESIGN AND SAMPLING. Plants were arranged in a which were subtended by bracts that formed an involucel and held completely randomized design. Three meristem samples were by a peduncle. Primary umbels consisted of two or more pe- removed from each cultivar at 17 collection times over an 85-d duncles, which together were subtended by an involucre. Floret period. Samples were taken at the following times: the day the plants number on an inflorescence ranged from one to 10. The first arrived at the University Park campus, 10 d later (plants had been inflorescence initiated on plants of ‘Duchess’ had an average of exposed to 100 mol of cumulative irradiance) and every 5 d [at 50 nine florets, while subsequent inflorescences on a plant had an mol photosynthetic photons (PP) intervals] thereafter. average of five florets. The average number of florets in the first Meristems were taken by removing 5 to 10 mm of tissue from inflorescences initiated on ‘Jennifer’ was seven. Subsequent the apical shoot tip. They were preserved in FAA (37.5% form- inflorescences had not developed fully in ‘Jennifer’ at the termi- aldehyde: 70% ethanol: 100% glacial acetic acid, 1:8:1 by vol.) nation of the experiment. fixative before examination with a stereo microscope using 25× FLORAL INITIATION. The vegetative meristem was convex with oculars and a 10× objective. Photographs were taken using leaf primordia developing in a spiral around the axis of the stem scanning electron microscopy. (Fig. 1A). At the onset of floral initiation, clefts became visible Fig. 1. Scanning electron micrographs showing Pelargonium ×domesticum apical meristem as it changes from the vegetative to the reproductive stage. (A) Dome shaped vegetative meristem (V) with two visible leaf (L) primordia. (B) Transitional meristem with five primordial meristems (PM) separated by clefts. A leaf primoridum (L) is visible in the lower left. (C) Shoot apex in which different meristem types are visible. The lower meristem is subtended by a leaf primordium (L), indicating a vegetative meristem (V). Other meristems lack leaf primordia, but are organized into inflorescences. These are subtended by bracts (B). (D) Three
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