The Chemical Induction of Parthenocarpy in the Calimyrna Fig and Its Physiological Significance' Julian C
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606 PLANT PhIYSIOLOGY 14. SCHREVEN-, D. A. \VANT. 1959. Effects of added 18. THORNTON, H. G. AND J. RuDIORF. 1936. The ab- sugars and nitrogen oni nodulationl of legumes. normal structure induced in nodules on lucerne Plant Soil 11: 93-112. (Medicago sativa L.) by the supply of sodium ni- 15. TANNER, J. W. AND I. C. ANDERSON. 1963. Ani trate to the host plant. Proc. Roy. Soc. Lod(lon external effect of inorganic nitrogen in root nio- B 120: 240-52. dulation. Nature 198: 303-04. 16. THIORNTON, H. G. 1936. The actionl of sodium ni- 19. TONHAZY, N. E. AND M. J. PEI.CZAR. JR. 1954. trate upoIn the inifectioii of lucernie root liairs by Oxidation of indoleacetic acid by anl extracellular nodule bacteria. Proc. Roy. Soc. London B 119: enzyme from Polyporus zersicolor anid a similar 474-92. oxidation catalyzed by nitrate. Science 120: 141- 17. THORNTON, H. G. The discovery of the use 42. of dichlorophenoxyacetic acid in controlling plant 20. WILSON, P. W. 1940. The biochemistry of symbio- growth. Rept. Rothamsted Expt. Sta. for 1939- tic nitrogen fixation. University of WViscoisin 1945: 83. Press, Madison. The Chemical Induction of Parthenocarpy in the Calimyrna Fig and its Physiological Significance' Julian C. Crane Department of Pomology, University of California, Davis, California Several auxins (1, 4, 5, 6) and a gibberellin (8) Materials and Methods have been found to be effective for inducing parthe- nocarpy in the Calimnvrna fig. None of the chemi- The investigation was coinducted at the Univer- cals, however, has been adopted by the industry for sity's Wolfskill Experimental Orchards, Winters, commercial application because of the inferior quality California, and the methods used were essentially of the figs produced. In contrast to pollinated fruits the same as those previously reported ( 1, 4). The that contain numerous drupelets with lignified endo- growth regulators were applied as aqueous solutions carp tissue which imparts a seedy texture, no ligni- on June 25, 1964, by spraying the young syconia and fication of this tissue occurs in chemiiicallv induced foliage to the point of slight drip. Each solution also parthenocarpic fruits, except those treate(d with ben- contained 0.05 % Tween 20 (polyoxyethylene sorbitan zothiazol-2-oxyacetic acid (6). The lercent parthe- monolaurate) as the surfactant. Tordon concentra- nocarpic fruit set with the latter chemical, however. tions of 20 mg/liter and below were tused, as severe is so erratic from year to year that it cannliot be relied leaf and wood damage resulted from higher concen- tupon. Conseqtuently, the search has continuied for a trations in 1963. SD 83393, a benzyladenine in which growth regulator that consistently indutces the produc- the hydrogen onl carbon atom 9 is substituted with the tion of large qtuanitities of parthenocarpic figs con- nonpolar ring struicture of tetrahydropyran, was tested taining drupelets with lignified endocarp. Prelimi- narv tests in 1(63 indlicated that the herbicide Tordon 2 (4-amino-3, 5, 6-trichloropicolinic acid) offerecl con- 2 Supplied by the Dow Chemical Company. Tordon is siderable promqise in this respect. The results ob- classified here as an auxin because of its similarity in biological activity to that of the substituted phenoxyacetic tained with this auxin in 1964, together with those acids. of a kinin compound are presented here. 36- (benzylamino) -9- (2-tetrahydropyranyl) -9H-purine, produced in the laboratories of Shell Development Com- pany and supplied by Dr. J. van Overbeek, Modesto, I Received November 27, 1964. California. CRANE-INDUCED PAtTHENOCARPY IN FIG 607 at concentrations of 500 and 1000 mg/liter. This though concentrations of 6 mg'/liter and below pro- new synthetic kinin was not only applied as a spray duced parthenocarpic fruit sets of only 88 to 94 %, but it was also injected with a hypodermic syringe these were considerably greater than those generally through the ostiole and into the central cavitv of the obtained by cross-pollination under commercial or- young fruit. The cavity was filled with 1 to 2 ml chard conditionis ( 1, 4, 5, 6). The percent fruit set of the solution. As a basis for comparison, GA, resulting from cross-pollination in the orchard where (potassium salt of gibberellic acid) and the auxin this investigation was conducted was not determined PCPA (para-chlorophenoxyacetic acid), regulators because the commercial procedure for pollination was proven to be effective for inducing parthenocarpy in not followed. the fig (1,8), were used at acid equivalent concen- Maturity of the parthenocarpic fruits, as judged trations of 25 and 75 mg/liter, respectively. Fifteen by color, firmness, and taste, occurred about a week to 20 fruits were used for each treatment. earlier than that of pollinated ones. Externallv, they At the time of treatment, the female flowers with- appeared identical to fruits that had been pollinated; in the syconia were receptive to polliniation. Cross- internally, however, the pulp was amber as contrasted pollination with the male fig, however, was prevented to the strawberry red that characterized pollinated by covering the branches bearing the fruits to be fruits. Ini contrast to parthenocarpic fruits pro- treated with muslin bags. The bags enclosed the duced with PCPA in which ligniification of the endo- branches for about 2 weeks, the period during which carp in the individual drupelets does niot occur, pollination of unbagged fruits was accomplished by this tissue in fruits produced with Tordon was the insect Blastophaga pseues. Bagged but untreated definitely lignified (fig 1). The degree of lignifi- fruits served as controls. Sinlce previous tests with cation, however, was not as great as that in pollinated the surfactant Tween 20 showed it to have no effect fruits, as (letermiined by mastication, but progressively on the fig, it was not applied in aqueous solution to increased as concentration of Tordon increased. the control fruits. Unless cross-pollination occurs The fruits had a seedy texture similar to pollinated or the syconia are treated with certain growth regu- ones; there was no evidence of embryos within the lators, they cease growth, wither and absciss in about individual drupelets. 2 weeks. No vegetative damage whatsoever was noted from concentrations of Tordon of 6 mg/liter or less. A Results concenitration of 8 mg/liter produced slight chlorosis of the younger leaves: leaf chlorosis and necrotic Tordon is a very effective compound for inducing areas on current-season's shoots increased progressive- parthenocarpy in the fig (table I). This chemical ly as concentration was increased above 8 mg/liter. at a concentration of 8 mg/liter was just as effective Slight leaf chlorosis developed following the applica- as 75 mg/liter of PCPA, the concentration of the tion of PCPA. This symptom did not occur in pre- latter that has been shown to be optimum (5). Al- vious experimentation with PCPA. perhaps because Table I. The Comiipairative Effectiveness of 2 Auixins, a Gibberellin, and a Kinin for Inducing Parthenocarpy in the Calimyrna Fig Parthenocarpic Treatment fruit set Observations on vegetative growth mg/liter Control (unpollinated) 0 Auxins: Tordonl 2 94 Like control 4 90 Like control 6 88 l ike control 8 100 Slight leaf chlorosis 10 93 Moderate leaf chlorosis and slight shoot injury 20 100 Severe leaf chlorosis and severe shoot injury PCPA 75 100 Slight leaf chlorosis Gibberellin: GA., 25 100 Rest period of 33 % of terminal vegetative buds was broken with subsequent shoot growth Kinin: SD 8339-sprayed 100 18 Abscission of youngest leaves 500 57 Abscission of youngest leaves SD 8339-inj ected 100 100 Like control 500 100 Like control 6,08 PLANT PHYSIOLOGY for inducing parthenocarpy lut, as shown in talble I, A the rest periodl of 33 % of the buds was broken anld ani average of 4.5 cm of new shoot growth resuilted. SI) 8339 was also very effective for inlduicinig )arthenocarpy, particularyv wheni inijecte(I inlto the younlg svcoinia (table 1). Iinjectioni of either coin- centration resuiltedl in 100 % p)arthenocarpic set and( (levelopmenit to maturity al)out a week earlier thani milatuirity of pollinate(l fruits. \Vhen applied as a 2 spray at conicenitrationis of 100( and(l 500 mg/liter, SD 8339 set onily 18 aln(d 57 % of the fruits, respectively. TI hese results inidicate that the compound is niot rea(Oi- 1v absorbed 1y fig leaves or that it nlax nlot be easily tralislocatedl. Externally, the fruits apleared as polli- nated onies, btit internally thev were similar t) frulits prodticedl with PCPA or GA. (fig 1). Abscission of the youn-est leaf on shoots that were sprayed with either conicenitratioll of SI) X33() ,3 d, J was the only vegetative response niote(l. Discussion \'ariotis auxiins, gibberellin-like, and kiinetinll-ike substances have been found to occtcr ill fruits: the seeds are partictilarly rich souirces of these materials (9). This informationi, together with the fact that .0~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..........'| X fruit size an(l shape are positively correlated in sev- .!.N.. ..4.g. eral inistances with see(d number anlCI distribution. le(d to the comiimonly accepted view\ that fruit grow\tll anld development are controlled 1yl hormilolnes emianat- *, ...Q inig fromii the see(ls. However, studies of auxinls in general have revealed positive correlationis between their levels and( the dlevelopmlient of thie eildospermi an(l embryo in the see(d btt little or nlo relationislhip) haS *e~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~. been shown between their leve's an(l fruiit gro\wth. Similarly, relatixrely large qtuanitities of gibberellin-like Fio. 1. 1C)Calumnvria figs by produced cross-pollina- substanices have been extracted from votinig see(ds. tioIn and those l)ro(duce(l by aqueous parthenlocarpically In general, a positive correlation wvas found(l between spray of 2) 6 apI)lication ing/liter of Tordon (4-amino-3, growth of the seed and inlcrease in amounllt of 5.