Controlling Height with Temperature Robert Berghage ‘Aoorrionat moex wonos, DIF, internode length, stem elongat iod, light, genetics, yield » chlorophyll, photope- ‘Summany. Temperature management has emerged as an important tool for plant height control in greenhouse production systems. This is particularly important in vegetable trans plant production where chemical controls for plant height are limited or not legal. Plant height is a function of the number of nodes and the length of each intemode, and both are strongly influenced by greenhouse temperatures. Node number, or formation rate, is primarily f function of the average greenhouse temperature, increasing as the average temperature jncreases. Internode length is strongly influenced by the relationship between the day and. night temperature, commonly referred to as DIF (day temperature ~ night temperature). As DIF increases, so does internode length in most plant species studied. Although the nature and magnitude of temperature effects vary with species, cultivar, and environmental conditions, these two basic responses can be used to modify transplant growth. Although data are limited, controlling transplant height with temperature does not appear to adversely influence plant establishment or subsequent yield. ontrolled-environment production of transplants, including vegetables, bedding plants, ornamentals, and forest trees, is one of the major commercial uses of greenhouse facilities in the United States. Transplant production systems have become increasingly auto- mated as new equipment has been developed in the latter half of the 21st century, but the successful application of this automation is largely depen- dent on growing high quality uniform transplants. Development and Widespread use of computer-controlled environmental management systems has allowed the investigation and implementation of new control strategies where the environment is modified to regulate the rate of crop development and the morphology of the plant to improve transplant quality. ‘Temperature is the one of the most easily and frequently modified environmental factors influencing plant growth. One can find many carly references documenting the profound effects of temperature on plant developmental rates, and morphology, Interactions between thermope- riod and photoperiod were well described as early as the 1940s and 1950s, (Went, 1944, 1957). This basic information remained a scientific curiosity, until the development of automated environmental control equipment made it possible to manage greenhouse temperature with a precision that was impractical before ‘Depameat of Horr, Pennnivnia Sate Univesity, Ua Pak PA TARO Heung - October December 1998.84) 535In the 1980s, a number of re search groups in the United States and Europe began to reexamine the effects ‘of temperature on the growth of plants in the greenhouse. The primary objec tive of these studies was t0 provide commercial producers with the infor- mation needed to take full advantage of the new control technologies (ic., Karlsson et al, 1983). During one of these studies, while examining the e fects of day and night temperatures between 14 and 30 °C on growth of Easter Lily (Lilinm longiflorum ‘Thunb.), it was observed that there ‘was an interaction between day and night temperature that affected stem length (Erwin et al., 1989). This re sponse was fit toa series of mathemati- cal models. It was discovered that the interaction could be described with a single term made up of the day tem- perature minus the night temperature. This approach was then applied to a number of other crops. including Went's (1944, 1957) tomato (Lyeo- persicon esculentum Mill.) data. It ‘quickly became apparent that this rela- tionship, coined DIF (Erwin et al., 1989), could be used to describe much ‘of the’ stem elongation response to diurnal thermoperiod and. photope- riod interactions (Erwin et al., 1989; Karlsson et al., 1989, Berghage and Heins, 1991), Subsequentresearch has shown near universality ofthe general response (Enwin and Heins, 1995), but it has also demonstrated that the magnitude, and nature of the thermomorphogenic effects varies be- tnween plant species, and among cult vvars within a species, as well as with timing and duration of the tempera ture fluctuations (Erwin and Heins, 1995; Myster and Moe, 1995; Vogelezang et al., 1992). ‘The manipulation of thermom- “onphogenic stem elongation responses has been broadly applied in commer- cial horticulture to reduce, or in some cases eliminate the use of chemical growth regulators (Roberts, 1991). This has been particularly importantin the production of transplants where concerns about chemical residues may make theiruse undesirable, ori many eases illegal Managing plant growth with temperature Pant height or stem length is simply the sum of the lengths of each of the internodes. It follows then, that 536 tomanage stem elongation, itisneces- sary to control or manage internode number, intemode length, or both Many environmental oF genetic fac tors, and endogenous and exogenous chemicals can influence either node number orinternode length. The pur- pose of this report is to review the effects of temperature on transplant height and quality. Other factors will be considered onlyas they relate to, or interact with temperature, responses. Node number ‘The rate of node development is, like most other developmentates(.e., Karlsson et al., 1989), driven prima” rily by shore and long term average temperature. Leaf unfolding rate (a measure of node, and intemode for- mation) shows a curvilinear response to temperature, increasing as tempera- ture increases to an optimum, then decreasingiftemperature becomiestoo high. For many plants optimum tem peratures for leaf unfolding fall be- tween 22 and 30 °C. For example, with sweet pepper (Capsicum annuum L.)scediings, Yapingand Heins(1996) reported maximum leaf count at an average temperature of =26 °C. Other greenhouse crops with optima in this Tange include poinsettia (Euphorbia pulcherima Willd. ex Klotzsch). (Berghage, 1989), dahlia (Dablia pinnatn Cav.) (Brondum and Heeins, 1993), and Boston fern ( Nephrolepis exaltata (L.) Schott) (Erwin et al., 1993), Somespecieshave optimaabove ‘or below this range. For example, op- imum temperature for leaf unfolding exceeds 30 °C in Easter lily (Karlsson etal, 1988), and 30 to 35 °Cin vinca [Catharanthusreseus(L.)] (Pietsch et al, 1995). In each ofthe studies cited above, average daily temperature was highly correlated with leaf unfolding While the relationship between day and night temperature (DIF) was not. While the rate of leaf unfolding may not be influenced by DIF, the total node number often is. This is fre- ‘quently reported in determinant spe- cies where flower initiation is influ- enced by night temperature (poinsct- tia, Berghage, 1989), or temperature fluctuations, [chrysanthemum (Den- dranthema grandiflora T2velev) Jensen, 1993}. ‘The rate of temperature-driven leaf unfolding in many plants is influ enced by light. Leaf unfolding rate increases as irradiance increases, (Pietsch et al., 1995) or photoperiod lengthens (Kozai et al., 1995), al: though much of this response is often attributed to increased leaf temper ture, Other factors may also influence the rate of node formation. For ex- ample, McCall and Atherton (1995) reporta lower leaf number fortomato plants grown with high salinity com- pared with low salinity in a positive DIF environment. Internode length Response rooir. Internode length in many plantsis greatly influenced by diurnal temperature fluctuations. Day— night temperature fluctuations (DIF) have been used to describe some, or in a few cases, nearly all of the tempera ture-driven internode, orstem clonga- tion responses in plants. For example, Easter Lily internode elongation (length) increases as DIF increases between -16 and 16 °C (Erwin et al., 1989). The response in lily is cunvilin carwith the magnitude of the response increasingasDIFincreases. In practice this means commercial growers can often achieve adequate height control by simply reducing a large positive DIF toa smaller positive or zero DIF. ‘The effect of DIF on internode length isduetoincreased ellelongation rather than an increase in cell number in Easter lily (Erwin et al., 1994). Stem and internode length responses to DIF have been reported for many other formamental and vegetable crops in- cluding: Dendranthema grandiflora (Karlsson et al., 1989b), Euphorbia pulcberrima (Berghage and. Heins, 1991), Xanthium pensylranicum L. (Erwin, 1991), Nephrolepis exaltata (Erwin’et al., 1993), Streptocarpus nabilisC.B, Clark Enwin, 1991), Cam ‘panula isophylla Moretti (Moe et al., 1991), Kalanchoe blosfeldiana Poelln (Mortensen, 1994), Besoniaxhiemalis Fotsch (Grindal and Moe, 1994), Be- goin xcheimantia TH. Everett ex ‘Web. (Bakken and Moe, 1995), Catbaranthus roreus (Piersch et al, 1995), Brassica (B. napus L. rapifera Metzg., B. oleracea L capitata L., and B, oleracea L. gemmifera Zenk.) (Bakken and Flones, 1995), Solanum suberosum L. (Kozai et al, 1995), Cucumis satius L. (Grimstad and Frimanslund, 1993), Ljcapersicon es- culentuyn (Grimstad, 1993), and Cap- scum annus (aping and Heins, 1996). For a more complete list see Myster and Moe (1995). Hodichnkay «October December 1998 8(4)Plantsrespondrapidlytoachange _ umol-mslongerphotoperiods(16 reduced in direct response to the mag- in DIF, altered growth rates can often _h),andanegative DIF, while thelong- nitude of the low temperature pulse be observed in as few as 24h (Erwin estinternodeswere produced withlow _(between-2and=10°C), Incontrastto and Heins, 1995), The response does irradiance (70 mol:m7s") and posi-_therelatively consistentresponse totem- not appear to have much of aresidual tive DIF. In general, the response to perature pulse immediately after dawn, effect; that is, plants respond to the DIF has been found to decrease as pulses at other times ofthe day or night current environmental regime with photoperiod increases (Erwin and have provided inconsistent results A litdlelagorlongtermcarryover(Bergh-_ Heins, 1995; Mysterand Moe, 1995). _ temperature pulse before dawn was not age, 1989). Although a response to Nutrient solution salinity effects effectivein salvia Salvia splendensE.) or DIF has been described for nearly all the magnitude of the response to DIF petunia (Enwin, 1991) but has b plants examined, exceptions include in tomato seedling stem elongation effective with Byjoniaxhiemalis(Grindal spring bulbsorephemerals(Erwinand (McCall and Atherton, 1995). Re- and Moe, 1994). A temperature pulse Heins, 1995), the magnitude and na- duced internode length in response to__ provided during the middle ofthe dark ture ofthe responseisaffectedby other a negative DIF was greater with lower period has generally proved to be least environmental conditions, timing of nutrient solution salinity (EC 3) than effective in reducing stem elongation the temperature fluctuations during high salinity (EC 15). Plants had re- (Jensen, 1994; Grindaland Moc, 1994; thephoto-skotoperiods,genetics,and duced y, when grown under positive Envin and Heins, 1993). plant maturity, or growth stage DiEenvironment,however, since there Daily stem elongationina number EnvIRONMENTALFACTORSTHATCON- — were nO significant interactions be- of greenhouse crops follows a pattern FOUND On MODIFY THE DIF RESPOKSE.. tween water relations and DIF, the similar to the response to a low tem Daily average temperature influences authorsconcludethatthemechanisms perature pulse. Stem clongation is intemodelengthandthustheresponse for inhibitory effects may be different. greatest during the end of the night to DIF in many plants. Sweet pepper Generics. The magnitude and and beginning of the day, decreasing seedling internode length was corre- nature of the DIF response is influ- during the day and increasing. again fated with average temperature aswell enced by plant species and cultivar. during the night (Erwin and Heins, aswith DIF (Yapingand Heins, 1996). Easter lily response is fairly large and 1995; Lechamny et al., 1985). How- Poinsettia has an optimum average straightforword (Erwin ct al., 1989). ever, thispattern does not appear to be temperature for internode elongation In contrast, plant height and intemn- consistent for all plants and the rela~ of=24°C, aswell asa correlation with ode length in Kalanchoe blosfeldiana tionship doesnot seemcorrelated with DIF (Berghage, 1989). Similarly, increases with either positive or nega- response totemperature DIF orpulscs, Grimstad and Frimanslund (1993) re: tive DIF (Mortensen, 1994; Jensen, For example, Bertram and Karlsen port independent effects of average 1994). In a trial of 20 seed geranium (1994) report that there was no clear temperature, and DIF on cucumber (Pelargonium xbortorumBailey) cukti- pattern in elongation in the DIF-sen- internode length. Internode length vars, Strefeler (1995) found that, al- _ sitiveplantspoinsettia or pelargonium, increased with increasingaverage tem- though the potential for elongation and that in petunia, also a DIE-sensi peratureand increasing DIF,cxceptat varied among cultivars, the response _tivespecies, elongation occurred mostly low day temperature (15 °C) where to DIF was the same. Euphorbin pul- during the day there was little response to DIF. cherrima cultivars however, vary in EFFECTS OF DIF ON OTHER QUALITY Tnternodeclongation responseto their response to DIF (Moc et al., FACTORS FOR TRANSPLANTS. Lcaf expan DIF is affected by light and photope- _1992a, 1992b). Tomato, corn, (Zea _sionand orientation can be affected by riod. Light quality has been shown to. mays L.) and cucumber have strong DIF (Enwin and Heins, 1995). Leat influence the response to DIE, pre- responses to DIF, while squash, (Cu area of Solanisms tuberosum plantlets sumably through effects related to curbita melopepo var. pepo C. water- (Koziietal., 1995), and Brassica trans- phytochrome photocquilibria (Moe — melon, (Citrudlus/anatus Thunb. pea plants (Bakken and Flones, 1995) was and Heins, 1990; Myster and Moe, (Pisum sativum L,),and bean (Phaseo- reduced when grown with a negative 1995;Enwinand Heins, 1995).Inean- fs ulgarisL.) are less responsive (Er- DIF. Likewise, Yaping. and Heins descent lighting used for day length win and Heins, 1995). (1996) report that leaf arca of sweet extension can eliminate a plant’s re- ‘Tiina OF TEMPERATURE CHANGES. —_pepperseedlings was highly correlated sponse to negative DIF while luores- ‘Thetiming and length of temperature with DIF, but in this case, only when entlightingcan enhancetheresponse changes influences the response to average daily temperature was consid- (Moc et al, 1991), Increasing ieradi-_ photo-skotoperiod temperature re- ered, In contrast, leaf area of Easter lily ance decreases DIF response (Myster sponses. Although there area number __ was correlated with night temperature and Moe, 1995). For example, in- of conflicting reports in the literature and not DIF (Enwin ct al., 1989) creasing irradiance from 100 to 200 (Erwin and Heins, 1995), there is a Leaves on plants grown with a nega himol-m2.c! ina growth chamber de- general consensusthatmany plantsare tive DIF tend to point downward, Creased thestem clongation ofpetunia_sensitivetoa temperature drop,orlow while those grown with a positive DIF (Petunia xinbrida Hort.) in response temperature pulse, during thefirst2to point upward (Enwin ct al,, 1989). to a positive DIF compared with 2 4h of the photoperiod (Myster and Leaf chlorophyll content is re- negative DIF (Kaczperskietal.,1991), Moe, 1995). For example, alow tem- duced in plants grown with a negative Kozai et al. (1995) report that with perature pulse given during the first 2. DIF (Berghageetal., 1991). Total leaf Solanum tuberosum, the shortest in- _h of the day reduced internode length chlorophyll increases as DIF increases. temodes were produced with plants of cucumber and tomato seedlings in Fuchsia (Fuchsia xbybrida Hort. ex ‘grown with higher irradiance (140 (Grimstad, 1993). Plant height was Vilm.)and Dendranthema Erwinand Hldichnolgy = Osober-Decemiber 1998 84) 537Heins ,1995). Reduced leaf chloro: phyil results in visibly chlorotic plants in negative DIF environments. For example, sweet pepper leaf reflectance at 580 nm decreased as DIF increased (Yaping and. Heins, 1996). DIE-in- duced leaf chlorosisis often reversible, with plants greening rapidly after re: moval from the negative DIF environ ment (Erwin and Heins, 1995). Lower day temperatures and re duced leaf chlorophyll have been con- dered responsible for the frequently reported reduction in dry weight of plants grown in negative DIF environ: ints, Grimstad (1993) reported re duced plant dry weight in both cucum ber and tomato in response to a low temperature pulse. Likewise, dry matter production in chrysanthemum is re- duced in negative DIF grown plants (Jensen, 1993). Negative DIF has been shown to reduce total soluble carbohy- drate content of Lilinm (Miller etal. 1993), and affects carbohydrate part. tioninginmanyplants(Enwinand Heins, 1995). In contrast, there are a number ofreports where DIF did not influence plant dry weight (Korii et al., 1995; Bakken and Flones, 1995), or where plants grown with a negative DIF had ‘greater dry weight than those grown \itha positive DIF (Bakken and Flones, 1995), ErrEcts oF Dif ON TRANSPLANT ES- ‘TaBLISHMENT AND YieLD. Although little research has been reported on the ef fects of DIF on transplant establish: ment and yield, work to date suggests that using a negative DIF to control transplant height does not adverscly affect establishment orcrop yields, and may in some cases improve yield or ‘crop quality. Bakken and Flones(1995) report that swede (Brassica napus L. rapifera Metzg.) dry weight at harvest .was not affected by transplant produc- tion temperature regime, but that har- ested root quality was better fiom transplants grown with anegative DIF. ‘They also reported that yield of cab bage (Brassica oleracea L capirata L.) and Brussels sprouts Brasica oleracea L. gemmifera Zenk.) was increased using negative DIF-grown transplants. Similarly, early yield and fruit quality of greenhouse cucumber was unaf- scted by alow temperature (Grimstad, 1993). Early fruiting in greenhouse tomato was delayed bya ow temperature pulse, but only when given for long time periods (4 or 5 weeks) and large temperature differ- 538 cences (9 and 6 °C) (Grimstad, 1993) Adequate height reduction could be obtained with a shorter treatment pe riod orsmallerpulse Grimstad, 1993) Conclusion Managing plant height in the greenhouse is an important quality consideration for transplant produe crs. Alternatives to the use of chemical growth regulators are needed because of environmental, or health concerns, and the unavailability of chemical con- trols for many plants. Height control through temperature management is achieved by two basic mechanisms, reduced node and internode number, primarily a function of average tem? perature, and reduced internode elon gation, primarily through increased night and or decreased day tempera tures, or low temperature pulses at or near the beginning of the day (nega tive or zero DIE), Although negative DIF has been shown to reduce leat chlorophyll, and plant dry weights in some cases, this has not translated into, adverse effects on transplant establ ment or yield. The most effective man- agement strategies will ake both num- berand internode length into account to produce compact transplants using, greenhouse temperature as one of a number of height management tools. Literature cited Berghage, RD. 1989. Modeling stem clon: gation in Euphorbia pulcherrima. PhD dis, ‘Michigan State Univ., Fast Lansing, Berghage, RD., JE. Erwin, and RD. Heins. 1991. Photoperiod influences leaf chlorophyll content in chrysanthemum, {grown with a negative DIF temperature regime, HortScience 26:92, Berghage, RD. and R.D. Heins. 1991. ‘Modeling poinsettia stem elongation, J ‘Amer. Soc. Hort. Sei. 116(1):14-18, Bakken, A.K. and M. Flones, 1995, Mor: phology:and field performance of Brassica transplants propagated under different day and night temperature regimes. Scientia, Horr, 61:167-176, Bakken, AK. and R. Moe. 1995, Height and quality control in Christmas begonia by growth-retarding remperature regimes, Acta Agr. Scand, Sect. B. Soiland Plant Sc 45:283-292, Bertram, L. and P. Karlsen. 1994. A com: parison sudy'on stem elongation of several greenhouse plants. Scientia Hort. 59:2 274, Brondum, J.J. and RD. Heins. 1993, Modeling’ temperature and photoperiod ‘effects on growth and development of dahlia. J, Amer. Soe. Hort. Sc. 118(1):36- 2. Enwin, LE., RD. Heins, and M.G. Karls son, 1989. "Thermomorphogenesis in Li inom longifiarum Thun. Amer. J. Bot. 7647-53, Enwin, .E. 1991. Thermomorphogenesis inplants. PRD diss, Michigan State Univ., East Lansing, Envin, JE, RD. Heins, and J.B, Faust 1993. Thermomorphogenicandphotope riodic responses of Nephrolepis exaltara ‘Dallas Jewel”. HonScience 28(3):182— 184 Enwin, LE, P. Velguth, and R.D. Heins 1994 Day /night temperature environment affects cell elongation but not division in Lilian longiflorum Thanks, J. Expt. Bot, 45(276):1019- 1028, Enwin, LE. and R.D. Heins. 1995. ‘Thermomorphogenic responses in stem and leaf development. HortScience 30(5):940-949, Grimstad, S,0. 1993. The effect ofa daily low temperature pulse on growth and de velopment of greenhouse cucumber and tomato plants during propagation. Sciea 353-62 Grimstad,S.0.and E, Frimanslund. 1993, lect of different day and night tempera ture regimes on greenhouse cucumber young plant production, flower bud for Imation and ca 53:191-204 Grindal, G. and R. Moe. 1994, Eficcrs of femperature-drop and a short dark inter: ruption on stem elongation and flowering in BegoniaxdiemalisFotsch Scientia Hon. 57:123-132, Jensen, HLE.K. 1993. Influence of drs tion and placement of a high night tem perature on morphogenesis of Dendran- ‘hema grandiflora Tzvelev, Scientia Hort 54:327-335, Jensen, HLE.K. 1994, Eficts of duration and degree of pulse-DIF temperatures on plant hejght and flowering of Kalanchoe Hlosfeldiana v. Poclin. Scientia Hor. 50:45-54 Kaezperski, M.P.,W.H, Caron, and M.G. Karisson. 1991. Growth and development Of Petunia xlpirida asa funtion oF tem perature andirradiance. J. Amer. Soc. Hor. Sci, 116(2):232-237. Karlson, M.,R.D. Hens, and W-H. Carl: son, 1983. Development of environmen tal strategies based on plant growth mod els. Acta Hort 147:153-160, Bofbctecg - acter Deeemter 1998.84Karlsson, M.G., RD. Heins, and J.B. Er win, 1988, Quantifying temperature con trolied leaf unfolding rate in ‘Nelle White™ Easter lily. J. Amer. Soc. Hort. Sci 113(1):7-74. Karlson, M.G., RD, Heins, LE, Erwin, and RLD, Benghage. 1989a. Development rate daring four phases ofchrysanthemum ‘growth as determined by preceding and prevallingtemperatures.J. Amer. Soc. Hor Sei, 114(2):158-162. Karlson, M.G., RD. Heins, JE. Erwin, ROD, Berghage, W.H. Carlson, and [.A. Bicmbaum. 1989b, Temperstureand pho: tosynthetic photon fhuxiniluence cheysan themum shoot development and flower fnitiation under shore day conditions. J ‘Amer, Soe, Hort, Sci. 114:158-163 Kovai,T., K, Watanabe, and B.R. Jeong. 1995. Stem elongation and growth of Solanum suberone 1. in vteo in response to photosynthetic photon flux, photope Fiod and difference in photoperiod and dark period temperatures. Scientia Hort. 64:1-9. Lecharny, A., H. Scwall, and E. Wagner, 1995, Siem extension rate in light grown, plants, Plant Phys. 79:625-629. McCall, D. and J.G. Atherton. 1995, In teractions between diumal temperature fluctuations and. salinity on expansion growth and water status of young tomato plants. Ana, Appl. Biol, 127:191-200. HerBatmnkgy + Ocier-Desemiber 1998 8(4) Miller, W.B.,P-A. Hammer,and TL. Kirk 1993, Reversed greenhouse temperatures alter carbohydrate status in Lilo Jona flsrum Thunb. ‘Nellie White’. J. Amer. Soc. Hort. Sci. 118(6):736-740, ‘Moe, R.and R-D. Heins. 1990. Control of plant morphogenesisand flowering by light Quality and temperature. Acta Hort, 272:81-89. Moe, R, RD. Heins, and JE, Ervin, 1991, Effect of day night temperature alterations, and plant growth regulatorson stem elongation and lowering ofthe long day plant Campanula ssoplylla Moretti Scientia Hort 48:141-181 Moe, R., T. Field, and LLM. Mortensen, 19923, Stem elongation and keeping qual ity in poinsettia (Enphorbin pulcberrina Willd.) as affected by temperature and supplementary lighting. Scientia Hort 50:127-130. Moe, R.,N. Glomsrud, I Bratberg, and S. ‘Vako, 1992b. Control of plant height in poinsettia by temperature drop and graphi- ‘al racking, Acta Hort. 327:41-48. Myster, J. and R. Moe. 1995. Effect of cliural temperature alternations on plant ‘morphology in some greenhouse crops a ‘mini-review, Scientia Hor. 62:205-215. Mortensen, L.M, 1994, Effects of day/ night temperature variations on growth, morphogenesis and flowering of Kalan: choc Mosféldiann v, Poelin. at different CO, concentrations, daylengths and pho- ton flux densities. Scientia Hort. 59:233- 241, Pietsch, G.M.,W.H, Carlson, RD. Heins, and -E. Faust. 1995. The effectof day and hight temperature and jiradiance on devel- ‘opment of Catharanshus aseus(L) ‘Grape Cooler’. J. Amer. Soc. Hort. Sci 120(5):877-881. Roberts, D.R. 1991. DIF. Greenhouse Mgr. 10(13):52-55. Strefler, MS. 1995. Growth response of 20 seed geranium cultivars to three day~ hight temperature regimes. HortScience 30(7):1465-1466, J. Vogelzang, R. Moe, H. Schussler, 1, “Hendriks, L. Cujpers,and E. Ueber. 1992, Cooperative European research on tem: perature strategies forbedding plants. Acta Hort. 327: 11-16. Went, FAW. 1984, Plant growth under controlled conditions. Il. Thermoperiod- icity in growth and fruiting of tomato, Amer J. Bot, 31:135-150. Went, EW. 1957. The experimental con- trol of plant growth. Chron, Bot. 17:1- 126. Yaping, S. and R.D. Heins. 1996. Inf cence of day and night temperatures on ‘sweet pepper seedling development. J Amer, Soc. Hort. Sci. 121(4):699-706. 39,