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The Control of the Annual Growth Cyclesof Six Temperate Fruit CropsJames G. Carew a & Nick H. Battey aa The Soft Fruit Technology Group, Centre for Horticulture andLandscape , The University of Reading , Whiteknights, Reading , RG66AS , UKPublished online: 12 Oct 2008.
To cite this article: James G. Carew & Nick H. Battey (2005) The Control of the Annual GrowthCycles of Six Temperate Fruit Crops, International Journal of Fruit Science, 5:1, 3-15, DOI: 10.1300/J492v05n01_02
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The Control of the Annual Growth Cyclesof Six Temperate Fruit Crops
James G. CarewNick H. Battey
ABSTRACT. Temperate perennial fruit crops typically show a clearlydefined cycle of growth through which they pass each year. In most, thiscycle is completed over two years so that to understand how growth iscontrolled, both years must be considered. Some, however, complete afull cycle of growth within one year. The objective of this paper is topresent the cycle of growth for four species: strawberry, raspberry, grapeand apple. Each species behaves differently in detail but there are, never-theless, some significant common features that provide opportunities fora greater understanding of underlying physiological mechanisms. [Arti-cle copies available for a fee from The Haworth Document Delivery Service:1-800-HAWORTH. E-mail address: <[email protected]> Website:<http://www.HaworthPress.com> 2005 by The Haworth Press, Inc. All rights re-served.]
KEYWORDS. Junebearing strawberry, everbearing strawberry, spring-bearing strawberry, primocane fruiting raspberry, biennial fruiting rasp-berry, grape, apple
James G. Carew and Nick H. Battey are Researchers with the Soft Fruit TechnologyGroup, Centre for Horticulture and Landscape, The University of Reading, Whiteknights,Reading, RG6 6AS, UK.
The authors would like to acknowledge funding from AMS Ltd., FAST Ltd., TheUniversity of Reading, DEFRA and HDC.
International Journal of Fruit Science, Vol. 5(1) 2005Available online at http://www.haworthpress.com/web/IJFS
2005 by The Haworth Press, Inc. All rights reserved.doi:10.1300/J492v05n01_02 3
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INTRODUCTION
The view has been expressed that the physiology of other Rosaceousfruit crops is not relevant to strawberry. The purpose of this paper is todemonstrate that there is a great deal to be gained by taking a broaderview.
BIENNIAL FRUITING RASPBERRY
The biennial fruiting raspberry has a clearly defined growth cyclewhich lasts two years, with vegetative growth taking place in the firstyear and flowering and fruiting in the second year (Figure 1). The cyclestarts in spring with growth of new shoots from either root buds oraxillary buds at the base of the main shoot (Hudson, 1959). Under fa-vorable environmental conditions the shoot passes through a phase ofcontinuous vegetative growth during which the canes develop leavesand internodes (Jennings and Dale, 1982). This continues until the au-
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BIENNIAL FRUITINGRASPBERRY
FIGURE 1. The Cycle of Growth of the Biennial Fruiting Raspberry.
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tumn when growth ceases and the cane forms a rosette of leaves at theapex of the shoot (Williams, 1959a), either because of the onset of dor-mancy or the initiation of flowers by the apical meristem (Moore andCaldwell, 1985). Flower initiation and dormancy are triggered by acombination of low temperatures and short photoperiods (Williams,1959b; Williams, 1960). During autumn and early winter, flower initia-tion continues but dormancy prevents flowers from emerging from thebuds and suppresses vegetative growth, dormancy being deepest duringNovember and December (White et al., 1998). These effects of dor-mancy are nullified through the action of cold temperatures or ‘chill-ing,’ six weeks at 3.3°C generally being sufficient to allow buds to growout when temperatures rise in the spring (Williams, 1959b). After this,the lateral shoots develop and bear the flowers initiated during theprevious autumn which then go on to produce fruit.
PRIMOCANE FRUITING RASPBERRY
In primocane fruiting cultivars (Figure 2), early cane growth occursin a similar way to biennial fruiting cultivars. Growth is affected by thechilling received by the root buds during winter (Carew et al., 2001).Growth continues throughout spring and into summer at a rate that isprincipally determined by temperature (Carew et al., 2003) untilAugust when flower initiation begins (Robertson, 1957). Increasingtemperature to 22-25°C, increased light intensity and an intermediatephotoperiod will all advance the start of flower initiation significantly(Carew et al., 2003; Hoover et al., 1989; Privé et al., 1993). Flower initi-ation happens earlier than for the biennial fruiting raspberry [because ofa difference in photoperiod and temperature sensitivity (Carew et al.,2000)] and so the primocanes are still actively elongating and have notyet entered dormancy. As a consequence, the flowers are able to growout immediately, during late summer and early autumn. Fruiting fol-lows and continues until the first frosts. In conventional production, thecanes are then pruned to the base to allow a new flush of canes to formthe following spring.
JUNEBEARING (SPRINGBEARING) STRAWBERRY
The growth cycle of the strawberry (Figure 3) has perhaps been mostcomprehensively studied over the years and so is the best understood. In
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September, flower induction is triggered by a combination of photoperi-ods less than 15 h and temperatures below 15°C (Guttridge, 1985; Itoand Saito, 1962; Sonsteby and Nes, 1998). Following this, flowers areinitiated at a rate that is determined by temperature, the rate increasingup to about 18°C and declining thereafter (Le Mière et al., 1996) but theflowers don’t grow out immediately, remaining within the apical bundleof leaf and stipule primordia until the following spring. At about thesame time as flowers start to be initiated in the autumn, the plants enterdormancy, a response greatly enhanced by a shortening photoperiodand lowering temperatures (Battey et al., 1998). Dormancy lasts forabout 4-5 months. Plants which do not receive chilling continue togrow, albeit at a reduced rate. About 8 weeks chilling at about 2°C isneeded for dormancy to be completed (Battey et al., 1998) although theamount of chilling required does vary depending on the measure used(Lieten, 1997). The cessation of dormancy combined with the highertemperatures of spring, allow elongation growth of flowers, which
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FIGURE 2. The Cycle of Growth of the Primocane Fruiting Raspberry.
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eventually begin to form fruit, fruiting lasting for about 4-6 weeks. Atthis stage temperature has a significant effect on growth. Higher tem-peratures cause rates of flower emergence to increase thereby advanc-ing fruiting, but partly as a result of this, berry size and yield decline (LeMière et al., 1998). Towards the end of fruiting, in mid-summer, theemergence of stolons from the crowns begins in response to lengtheningphotoperiods and increasing temperatures, signalling the start of runnerproduction (Durner et al., 1984).
EVERBEARING STRAWBERRY
The control of growth of the everbearing strawberry (Figure 4) hasnot been documented as widely as that of the Junebearing strawberry.Work is needed at a fairly basic level to correct this lack of information.That fruiting of everbearers occurs in more than a single peak of produc-tion (Perez de Camacaro et al., 2002) forms the main difference from
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FIGURE 3. The Cycle of Growth of the Junebearing Strawberry.
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Junebearing strawberries (Taylor, 2002). The first peak of croppingis during spring and early summer at about the same time as June-bearing cultivars and the second during late summer and autumn.This therefore suggests a fundamental difference in the control offlower initiation. Both Smeets (1980) and Durner et al. (1984)present data showing earliest flower initiation at temperatures be-tween 20 and 26°C and Dennis et al. (1970) have shown that longerphotoperiods cause earlier flower initiation. However, the two crop-ping peaks can overlap, suggesting that flower initiation doesn’t oc-cur in two distinct periods, controlled by temperature and photo-period, but throughout the year, except when growth stops duringwinter. Because of this, the timing of flower initiation shown in Fig-ure 4 is somewhat speculative.
Very little work has been published on dormancy in everbearers.Smeets (1980) found increased runnering with chilling in the twoeverbearers ‘Rabunda’ and ‘Ostara’ suggesting that everbearing straw-berries do enter dormancy in the autumn. From data collected over twoyears at Reading, however, it appears that dormancy in everbearers is
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FIGURE 4. The Cycle of Growth of the Everbearing Strawberry.
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much shallower than in Junebearers, being removed by a lower level ofchilling than would be required for Junebearers (Carew and Battey,unpublished).
APPLE
The growth cycle of the apple (Figure 5) occurs over two years. Itstarts with flower initiation in mid-summer, 2-3 months after fullbloom of the flowers initiated the previous year. This is a time whenfruit growth will be rapid (Bertelsen et al., 2002). For the other spe-cies described here, a degree of environmental control of flower ini-tiation has been shown but this is not the case for apple; indeed,Thomas and Vince-Prue (1997) classify the apple as day-neutral.Convincing evidence for control by the translocation of plant hor-mones has been presented over the years and related to biennialbearing (Buban and Faust, 1982). Gibberellic acid released from de-veloping seeds is believed to inhibit flower initiation. An antagonis-tic relationship between flower initiation and vegetative growth has
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FIGURE 5. The Cycle of Growth of the Apple.
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also been described (Forshey and Elfving, 1989) whereby vigorousgrowth tends to reduce flower initiation. Flower initiation lasts forabout 1-2 months (McArtney et al., 2001) and usually by the end ofthis period, vegetative growth of the trees has finished (Forshey andElfving, 1989). The plants enter dormancy in the autumn but again incontrast to the other crop species under discussion here, the apple is rel-atively insensitive to photoperiod at this time (Thomas and Vince-Prue,1997; Wareing, 1969). Apple does however require a period of chillingbefore growth is possible in the spring (Jacobs et al., 2002). Flowers ini-tiated during the previous summer emerge from buds at a rate which isdetermined by environmental conditions (Dennis, 1986). Fruiting fol-lows.
GRAPEVINE
Morphological development of flowers in grapevine is unique. Itis also particularly interesting because flowering is both environ-mentally and endogenously controlled (Buttrose, 1974). The grapeis therefore intermediate between the extremes of the apple, whereflowering is almost entirely controlled endogenously, and the June-bearing strawberry and the raspberry where flowering is more influ-enced by temperature, light intensity and photoperiod. This leads to agrowth cycle with elements in common with those of other fruitcrops (Figure 6).
Bud growth starts in spring with a burst of vegetative growth, re-sulting in the production of lateral shoots, in the axil of which form la-tent buds (Srinvasan and Mullins, 1981). The meristem within thesebuds may produce up to 10 leaf primordia until induced to produce ananlage, usually during early summer (Lavee, 1985). Anlagen are un-committed primordia (Srinvasan and Mullins, 1981), and can eventu-ally be induced to flower or to produce tendril primordia (Buttrose,1974), thereby acting as an intermediate step between a meristemforming leaf primordia and one initiating flowers or tendrils. Whetherinflorescence primordia or tendril primordia develop depends on envi-ronmental and endogenous conditions during summer. Increasinglight intensity and temperature to about 30/35°C causes inflorescencedevelopment to be prioritized over tendril formation (Buttrose, 1969).However, inflorescence formation is also markedly affected by thehormones cytokinin and gibberellic acid, cytokinin causing a shift to-
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wards inflorescence development, gibberellic acid towards tendril ini-tiation (Srinvasan and Mullins, 1978; Srinvasan and Mullins, 1980).The significance of this becomes apparent when it is remembered thatin addition to initiating new inflorescence primordia during summer,fruit will also be forming which may therefore affect hormonal trans-location within the shoot. During autumn, the plants enter dormancyin response to shortening photoperiods (Wake and Fennel, 2000) withone to four weeks of chilling at 0°C being required for its completion(Dokoozlian et al., 1995; Dokoozlian, 1999). Then, in the spring, justprior to and during bud burst, flower initiation occurs within the in-florescence primordia formed the previous summer (Srinvasan andMullins, 1981). The flowers continue developing throughout spring,emerging from the buds to form fruit during the summer. Two importantaspects of grape flowering are therefore that inflorescence initiationhappens at the same time as fruiting, during summer and the flowersdo not over winter, they are initiated in the spring of the year in whichthey form fruit.
James G. Carew and Nick H. Battey 11
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CYCLE OFGROWTH OF
GRAPE
FIGURE 6. The Cycle of Growth of the Grape.
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CONCLUSIONS AND FUTURE AREAS FOR STUDY
1. The distinction between primocane and biennial raspberries is auseful way of classifying cultivars. However, on further examina-tion, the distinction becomes less clear as primocane raspberriesshow characteristics analogous to the biennial raspberry and viceversa (Carew et al., 2000). Some biennial fruiting cultivars, suchas ‘Glen Moy’, initiate flowers sufficiently early to allow the pro-duction of a small amount of fruit at the tips of canes during theautumn in their first year of development (Ourecky, 1976), andare therefore somewhere between primocane and biennial fruit-ing. If not cut down at the end of the year, primocane fruitingcultivars can produce a second crop of fruit in the second year ofgrowth from the lower region of the cane, which fails to produce acrop in the first year. In this case, half the cane is primocane fruit-ing and half biennial fruiting. This has commercial significancebecause it allows double cropping of primocane fruiting raspber-ries (Jennings, 1996). Perhaps a better way of describing rasp-berry cultivars would be as part of a continuum based on theproportion of crop formed in the first or second year of growth.
2. The similarities between the Junebearing strawberry and the bien-nial fruiting raspberry are clear. Both require short days and lowtemperatures for flower induction, both require a period of winterchilling and both are similarly affected by spring and summertemperatures. Whether the everbearing strawberry and the primocanefruiting raspberry are also similar is not yet clear. What is neededis a comparison of the control of flower initiation in these twotypes of fruit crop.
3. In both apple and grape, flower initiation is affected by the concur-rent fruiting. GAs and cytokinins have been implicated in this, ashas the affect of fruiting on carbohydrate partitioning. Of the re-maining crops reviewed here, the everbearing strawberry and theprimocane fruiting raspberry both initiate flowers at the same timeas fruiting. Whether GA or cytokinin would have the same effectin these crops as in grape and apple is a topic of interest for futureresearch.
4. It is striking that in apple no effect of the environment on the in-duction of flowering and of dormancy has been demonstrated.More detailed study of temperature and photoperiod effects mightprove worthwhile.
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REFERENCES
Battey, N.H., P. Le Mière, A. Tehranifer, C. Cecik, S. Taylor, K.J. Shrives, P. Hadley,A.J. Greenland, J. Darby, and M.J. Wilkinson. 1998. Genetic and environmentalcontrol of flowering in strawberry, pp. 111-131. In: K.E. Cockshull, D. Gray, G.B.Seymour, and B. Thomas, (eds.), Genetic and environmental manipulation of horti-cultural crops. CAB International, Wallingford.
Bertelsen, M.G., D.S. Tustin, and R.P. Waagepetersen. 2002. Effects of GA3 andGA4+7 on early bud development of apple. J. Hort. Sci. Biotechnology 77:83-90.
Buban, T. and M. Faust. 1982. Flower bud induction in apple trees: Internal control anddifferentiation. Hort. Rev. 4:174-203.
Buttrose, M.S. 1969. Fruitfulness in grapevines: Effects of light intensity and tempera-ture. Bot. Gaz. 130:160-173.
Buttrose, M.S. 1974. Climatic factors and fruitfulness in grapevines. Hort. Abstr.44:19-326.
Carew, J.G., T. Gillespie, J. White, H. Wainwright, R. Brennan, and N.H. Battey. 2000.The control of the annual growth cycle in raspberry. J. Hort. Sci. Biotechnology75:495-503.
Carew, J.G., K. Mahmood, J. Darby, P. Hadley, and N.H. Battey. 2001. The effects oflow temperatures on the vegetative growth and flowering of the primocane fruitingraspberry ‘Autumn Bliss’. J. Hort. Sci. Biotechnology 76:264-270.
Carew, J.G., K. Mahmood, J. Darby, P. Hadley, and N.H. Battey. 2005. The effect of tem-perature, photosynthetic photon flux density and photoperiod on the vegetative growthand flowering of Rubus idaeus L. ‘Autumn Bliss’. J. Amer. Soc. Hort. Sci. (accepted).
Dennis, F.G. 1986. Apple. In: Monselise, S.P. (ed.), CRC Handbook of fruit set and de-velopment. CRC Press, Boca Raton, USA.
Dennis, F.G., J. Lipecki, and C.L. Kiang. 1970. Effects of photoperiod and other fac-tors upon flowering and runner development of three strawberry cultivars. J. Amer.Soc. Hort. Sci. 95:750-754.
Dokoozlian, N.K., L.E. Williams, and R.A. Neja. 1995. Chilling exposure and hy-drogen cyanamide interact in breaking dormancy of grape buds. HortScience30:1244-1247.
Dokoozlian, N.K. 1999. Chilling temperature and duration interact on the budbreak of‘Perlette’ grapevine cuttings. HortScience 34:1054-1056.
Durner, E.F., J.A. Barden, D.G. Himelrick, and E.B. Poling. 1984. Photoperiod andtemperature effects on flower and runner development in day-neutral, junebearing,and everbearing strawberries. J. Amer. Soc. Hort. Sci. 109:396-400.
Forshey, C.G. and D.C. Elfving. 1989. The relationship between vegetative growth andfruiting in apple trees. Hort. Rev. 11:229-287.
Guttridge, C.G. 1985. Fragaria ananassa. In: Halevy, A. (ed.), CRC Handbook ofFlowering, Vol III. CRC Press, Boca Raton, USA.
Hoover E., J.J. Luby, D. Bedford, M. Pritts, E. Hanson, A. Dale, and H.A. Daubeny.1989. Temperature influence on harvest date and cane development in primocanefruiting red raspberries. Acta Hort. 262:297-303.
Hudson, J.P. 1959. Effects of environment on Rubus idaeus L. 1. Morphology and de-velopment of the raspberry plant. J. Hort. Sci. 34:163-169.
James G. Carew and Nick H. Battey 13
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nloa
ded
by [
UO
V U
nive
rsity
of
Ovi
edo]
at 0
0:40
16
Oct
ober
201
4
Ito, H. and T. Saito. 1962. Studies on the flower formation in the strawberry plant. I. Ef-fects of temperature and photoperiod on the flower formation. Tohoku J. Agr. Res.13:191-203.
Jacobs, J.N., G. Jacobs, and N.C. Cook. 2002. Chilling period influences the progres-sion of bud dormancy more than does chilling temperature in apple and pear shoots.J. Hort. Sci. Biotechnology 77:333-339.
Jennings, D.L. 1996. Variation in the cropping seasons of two raspberries bred byMedway Fruits. Proc. ADAS/HRI/EMRA Soft fruit conference 1996, 83-87.
Jennings, D.L. and A. Dale. 1982. Variation in the growth habit of red raspberries withparticular reference to cane height and node production. J. Hort. Sci. 57:197-204.
Lavee, S. 1985. Vitis vinifera. In: Halevy, A. (ed.), CRC Handbook of flowering, VolIII. CRC Press, Boca Raton, USA. pp. 456-471.
Le Miére, P., P. Hadley, J. Darby, and N.H. Battey. 1996. The effect of temperature andphotoperiod on the rate of flower initiation and the onset of dormancy in the straw-berry (Fragaria �ananassa Duch.). J. Hort. Sci. 71:361-371.
Le Miére, P., P. Hadley, J. Darby, and N.H. Battey. 1998. The effect of thermal envi-ronment, planting date and crown size on growth and development in Fragaria
ananassa Duch cv. Elsanta. J. Hort. Sci. Biotechnology 73: 786-795.Lieten, F. 1997. Effects of chilling and night-break treatment on greenhouse produc-
tion of ‘Elsanta’. Acta Hort. 439:633-637.McArtney, S.J., E.M. Hoover, P.M. Hirst, and I.R. Brooking. 2001. Seasonal variation
in the onset and duration of flower development in ‘Royal Gala’ apple buds. J. Hort.Sci. Biotechnology 76:536-540.
Moore, J.N. and J.D. Caldwell. 1985. Rubus. In: Halevy, A. (ed.), CRC Handbook offlowering, Vol. IV. CRC Press, Boca Raton, USA. pp. 226-238.
Ourecky, D.K. 1976. Fall-bearing red raspberries, their future and potential. Acta Hort.60:135-144.
Perez de Camacaro, M.E., G.J. Camacaro, P. Hadley, N.H. Battey, and J.G. Carew.2002. Pattern of growth and development of the strawberry cultivars Elsanta, Bo-lero, and Everest. J. Amer. Soc. Hort. Sci. 127:901-907.
Privé, J-P., J.A. Sullivan, J.T.A. Proctor, and O.B. Allen. 1993. Climate influencesvegetative and reproductive components of primocane fruiting red raspberry cultivars.J. Amer. Soc. Hort. Sci. 118:393-399.
Robertson, M. 1957. Further investigations of flower bud development in the genusRubus. J. Hort.Sci. 32:265-273.
Smeets, L. 1980. Effect of temperature and day-length on flower initiation and runnerformation in two everbearing strawberry cultivars. Scientia Hort. 12:19-26.
Sonsteby, A. and A. Nes. 1998. Short days and temperature effects on growth and flow-ering in strawberry (Fragaria ananassa Duch.). J. Hort. Sci. Biotechnology73:730-736.
Srinvasan, C. and M.G. Mullins. 1978. Control of flowering in the grapevine (Vitisvinifera L.). Plant Physiol. 61:127-130.
Srinvasan, C. and M.G. Mullins. 1980. Effects of temperature and growth regulators on for-mation of anlagen, tendrils and inflorescences in Vitis vinifera L. Ann. Bot. 45:439-446.
Srinvasan, C. and M.G. Mullins. 1981. Physiology of flowering in the grapevine, a review.Amer. J. Enol. Viticult. 32:47-63.
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by [
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rsity
of
Ovi
edo]
at 0
0:40
16
Oct
ober
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4
Taylor, D.R. 2002. The physiology of flowering in strawberry. Acta Hort. 567:245-251.
Thomas, B. and D. Vince-Prue. 1997. Photoperodism in Plants. Academic Press, London.Wake, C.M.F. and A. Fennell. 2000. Morphological, physiological and dormancy responses
of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109:203-210.Wareing, P.F. 1969. The control of bud dormancy in seed plants. In: Dormancy and
survival. Symposia of the Society for Experimental Biology XXIII, 241-262, Cam-bridge University Press, UK.
White, J.M., H. Wainwright, and C.R. Ireland. 1998. Interaction of endodormancy andparadormancy in the raspberry cultivar ‘Glen Moy’. Ann. Appl. Biol.132:487-495.
Williams, I.H. 1959a. Effects of environment on Rubus idaeus L. II. Field observationson the variety Malling Promise. J. Hort. Sci. 34:170-175.
Williams, I.H. 1959b. Effects of environment on Rubus idaeus L. III. Growth and dor-mancy of young shoots. J. Hort. Sci. 34:210-218.
Williams, I.H. 1960. Effects of environment on Rubus idaeus L. V. Dormancy andflowering of the mature shoot. J. Hort. Sci. 35:214-220.
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1. Fill out the form below and make sure that you type or write out clearly both the nameof the journal and your own name and address. Or send your request via e-mail [email protected] including in the subject line “Sample Copy Request” andthe title of this journal.
2. Make sure to include your name and complete postal mailing address as well as yourinstitutional/agency library name in the text of your e-mail.
[Please note: we cannot mail specific journal samples, such as the issue in which a specific article appears.Sample issues are provided with the hope that you might review a possible subscription/e-subscription withyour institution's librarian. There is no charge for an institution/campus-wide electronic subscriptionconcurrent with the archival print edition subscription.]
I will show this journal to our institutional or agency library for a possible subscription.Institution/Agency Library: ______________________________________________
Name: _____________________________________________________________
Institution: __________________________________________________________
Address: ___________________________________________________________
City: ____________________Return to: Sample Copy Department,The Haworth Press, Inc.,
10 Alice Street, Binghamton, NY 13904-1580
State: __________ Zip: ____________________
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