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544 Kurze Originalmitteilungen Die Natffr- wissenschaften Iokal an der Injektionsstelle schnell wachsende und bald zum Tode Iiihrende Tumoren, die siimtlieh histologisch yon Pro- fessor Dr. H. HAIVlPXRL, Bonn, als Spindelzellensarkome dia- gnostiziert wurden. Da subcutane Sarkome bet unseren Ratten spontan sehr selten vorkommen (unter 0,5 %) und alle Tnmoren streng an der Injektionsstelle auftraten, kann der Kausalzusammenhang als gesichert gelten. Die hohe Ausbeute yon 7 unter 12 Ratten bet einer mittleren Gesamtdosis yon 500 mg spricht fiir eine beachtliche carcinogene Wirksamkeit des Parafuchsins. Die Arbeiten wurden durch die Deutsche Forschungs- gemeinschaft erm6glicht. Laboratorium der Chirurgischen Universit~itshlinik, Fret- burg i. Br. H. DRUCKREY, H.A. NIEPER nnd H.W. Lo Eingegangen am 2. November 1956 i) SCHILLER, W.: Amer. J. Canc. 3t, 486 (1937). -- HARRIS, P.N.: Canc. Res. 7, 35 (t947). ~) WILLHmM, R., u. A.C. Joy: Cane. Res. 12, 308 (1952). 8) DRUCKREY, H., u. D. SCHMXHL: Naturwiss. 8, 215 (1955). ~) YOSmDA, T., T. SHIMANCHI U. C. KI.~: Gann 35, 272 (1941). EiniluB des Gibberellins auf die B1/itenbildung bei Kalanchoe bloflfeldiana Wie wir frtiher mitgeteilt haben~), kann man bet der Lang- Kurztagpflanze Bryophyllum und bet der Langtagpflanze Lapsana communis unter streng vegetativen Bedingungen Bltitenbildung durch Gibberellin ausl6sen. Schon wesentlich Irtiher als bet diesen Objekten haben wir mit der Gibberellin- behandlung bet der Kurztagpflanze Kalanchoe bloBfeldiana begonnen. Obgleich wir die Behandlungsweise auf3erordent- lich stark variiert haben, ist his hente, d.h. mehr als 5 Monate seit Beginn der Versuche, an keiner der mehr als 150 Ver- suchspflanzen eine F6rderung des B1/ihens zu beobachten. Im Gegenteil, es macht sich eine gewisse Hemmung geltend: die Bltiten werden an den Gibberellinpflanzen zwar etwa zum gleichen Zeitpunkt sichtbar wie an den dutch Kurztag indu- zierten Kontrollen, aber es kommen nur relativ wenige Bliiten zur Entwicklung, nnd nach einiger Zeit findet ein Riick- schlag zur vegetativen Phase start. Auch das L~ingenwachstum der Internodien, das bet nichtblfihenden Kalanchoepflanzen wesentlich stbxker ist als bet bliihenden, ist bet den Gibberel- linpflanzen stark gef6rdert, und die Blattsukknlenz, die bet bliihenden Pflanzen besonders hoeh ist, ist bet den Gibberellin- Exemplaren gering. Bet Kalanchoe werden die vegetativen Merkmale also durch Gibberellin gef6rdert, die reproduktiven aber gehemmt. Aus dieser g~gzlich andersartigen IReaktion der Kurztag- pflanze Kalanehoe gegenfiber unseren anderen Versuchsobjek- ten ergibt sich, dab das Gibberellin keinesfalls der allein ausschlaggebende Faktor fiir die Bliitenbildung sein kann Evgl. ~)]. Mit Unterstfitzung der Deutschen Forschungsgemein- schaft. Gdttingen, P/lanzenphysiologisches Institut der Universit~l t~. HARDXR und R. B/3~sow Eingegangen am 26. November 1956 t BONsow, R., u. R. HARDER: Naturwiss. a) 43, 479 (1956); b) 43, 527 (1956). fiibberellin and Flower Formation As reported previously1), gibberellin induces flower for- mation in noncold-treated biennial tfyoscyamus niger, thus enabling this plant to bypass its normal cold requirement. The experiments have meanwhile been extended to other cold- requiring plants and to a number of long- and short-day plants, all of them maintained under noninductive conditions. This note is to summarize the results. A fuller account will be pub- lished elsewhere. Cold-requiring plants. Some, but not all plants of a variety of carrot (Early French Forcing) formed flowers after 11/2 to 21/o months of daily applications of t0 [zg or more of gibberel- lina). In another variety (DANvER'S Half Long), a bolting response only has been observed so far, and the same is true of parsley (Hamburg) and turnips (Purple Top Globe). The treatment is being continued. In winter rye (PETKUS) stem elongation was greatly promoted, but no effect on flower initiation was to be detected (same leaf number to first flower and same developmental stage of the terminal shoot meristem in treated plants and controls)3). Long-day plants. A rapid flowering response, equalling the effect of long-day exposure, was obtained in Samolus parviflorus with daily applications of 2 ~g or 5 ~g of gibberellin. Flower buds became visible after about one month, open flowers appeared one week later (5 ~zg treatment). A similar response seems to occur in Crepis tectorum (the experiment is still in progress). In Silent armeria and in annual ttyoscyamus niger the response was much slower and so far partial (only part of the treated plants responding, similar to the Early French Forcing carrot). The minimum effective dose was 10 txg daily, and the first flower buds became visible after about 7 to 8 weeks in Silene and after f0 weeks in Hyoscyamus. In Hyoscyamus, the buds failed to develop further; however, continued development may be a matter of proper dosage and periods of treatment. In Petkus spring rye, the results were identical with those in nonvernalized winter rye. Short day plants. A four week long application of 1 to /0 b~g gibberellin daily greatly enhanced stem elongation in cocklebur (Xanthium saceharatum), but the plants remained strictly vegetative. An experiment with soybean (Biloxi), still in progress, indicates a similar behavior. Comments. It is evident that gibberellin is capable of inducing flower formation in a considerable number of plants and under strict noninductive conditions. However, this effect of gibberellin does not seem to be general. Promotion of flowering was not observed in rye (winter and spring) nor in short-day plants kept in long-day conditions, although the vegetative effects of gibberellin application were very pro- nounced. Where flower formation did occur in biennial and long-day plants, the response was variable: rapid and com- plete in some species, slow and possibly partial in others. This variability cannot be attributed to differences in photoperiodic requirement, since the sensitivity of Samolus and Crepis to long-day induction is not greater than that of Hyoscyamus and Silene. At present, to account for the variable effect of gib- berellin on flower formation two explanations appear most probable : 1. Gibberellin, or a substance of similar physiological activity, is not the sole regulator of flower formation, but participates in this process in conjunction with other factors. Thus, the factors necessary for the complete functioning of gibberellin in flower formation may be limiting in some plants, but not in others. 2. The gibberellins of different plants are not identical and not completely interchangeable, so that a given gibberellin will not be equally effective in all plants, even if gibberellin is the factor limiting flower formation in these plants. Whether the action of gibberellin in flower formation is direct or indirect, remains to be seen. If the plants which form flowers after gibberellin treatment are induced in the natural way (by a cold treatment or by long days), bolting and flower initiation occur almost simultaneously. In contrast, gibberellin treatment causes stem elongation to occur first, while flower initiation may be more or less delayed. Therefore it seems possible that the primary effect of gibberellin is initia- tion of stem elongation, and that further processes are set in motion in the elongating stem which in turn lead to flower formation. In this case, flower formation would be a secon- dary effect of gibberellin. This work was supported in part by a research grant (RG-3939) from the National Institutes of Health, U.S. Public Health Service. Department of Botany, University o/ California, Los Ange- les 2g, Cal. ANTON LANO Eingegangen am t6. November t956 t) LANE A.: Naturwiss. 43, 284 (t956). 3) Part of the experiments was done with a mixture of gibberel- lin A and gibberellie acid, kindly supplied by Dr. F.H. STODOLA, U. S. D. A. Northern Utilization Research Branch, Peoria, Ill., part with a preparation containing gibberellic acid only, obtained through the courteousy of Dr. CURT LEBEN, Lilly Research Labora- tories, Indianapolis, Ind. 8) Rye was treated either by applying a drop of gibberellin solu- tion to the base of the blade of the youngest fully expanded leaf, or by immersing a leaf blade, with the tip cut, into the solution. The other plants were treated by applying the solution to the apical region of the shoot. Most experiments were carried out by Mrs. PATRJCIA B. FINN. Herausgeber und verantwortlicher Redakteur: Professor Dr, Ernst Lamla. G6tfingen. -- Veraixtwortlich ffir den Anzeigenteih Gfinter HoItz, Berlin W 35, Reichpietschufer 20. -- Springer-Verlag, Berlin-Gbttingen-Heidelberg. -- Drnck der Universitiitsdrnckerei H. Sttixtz AG., Wfirzburg. -- Printed in Germany

Gibberellin and flower formation

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544 K u r z e O r i g i n a l m i t t e i l u n g e n Die Natffr- wissenschaften

Iokal a n der In jek t ionss te l l e schnel l w a c h s e n d e u n d ba ld z u m Tode I i ihrende T u m o r e n , die s i imtl ieh h is to logisch yon Pro- fessor Dr. H . HAIVlPXRL, Bonn, als Sp inde lze l lensarkome dia- gnos t iz ie r t wurden .

Da s u b c u t a n e Sa rkome bet unse r en R a t t e n s p o n t a n sehr se l ten v o r k o m m e n (unter 0,5 %) u n d alle T n m o r e n s t reng an der In jek t ionss te l l e au f t r a t en , k a n n der K a u s a l z u s a m m e n h a n g als ges icher t gelten. Die hohe A u s b e u t e yon 7 u n t e r 12 R a t t e n bet e iner m i t t l e r en Gesamt dos i s yon 500 m g spr ich t fiir eine beach t l i che ca rc inogene W i r k s a m k e i t des Pa ra fuchs ins .

Die Arbe i t en w u r d e n du rch die D e u t s c h e F o r s c h u n g s - g e m e i n s c h a f t e rm6gl icht .

Laboratorium der Chirurgischen Universit~itshlinik, Fret- burg i. Br.

H. D R U C K R E Y , H.A. NIEPER n n d H . W . Lo Eingegangen am 2. November 1956

i) SCHILLER, W.: Amer. J. Canc. 3t, 486 (1937). - - HARRIS, P.N.: Canc. Res. 7, 35 (t947).

~) WILLHmM, R., u. A.C. Joy : Cane. Res. 12, 308 (1952). 8) DRUCKREY, H., u. D. SCHMXHL: Naturwiss. 8, 215 (1955). ~) YOSmDA, T., T. SHIMANCHI U. C. KI.~: Gann 35, 272 (1941).

EiniluB des Gibberellins auf die B1/itenbildung bei Kalanchoe bloflfeldiana

Wie wir fr t iher mi tge te i l t haben~), k a n n m a n bet der Lang - K u r z t a g p f l a n z e B r y o p h y l l u m und bet der L a n g t a g p f l a n z e L a p s a n a c o m m u n i s u n t e r s t r eng v e g e t a t i v e n B e d i n g u n g e n Bl t i t enb i ldung du rch Gibberel l in aus l6sen. Schon wesen t l i ch Irtiher als bet d iesen Ob j ek t en h a b e n wir m i t der Gibberel l in- b e h a n d l u n g bet der K u r z t a g p f l a n z e K a l a n c h o e bloBfeldiana begonnen . Obgleich wir die B e h a n d l u n g s w e i s e auf3erordent- l ich s t a rk var i ie r t haben , is t his hen te , d .h . m e h r als 5 Mona t e sei t B e g i n n der Versuche, an ke iner der m e h r als 150 Ver- suchsp f l anzen eine F 6 r d e r u n g des B1/ihens zu beobach t en . I m Gegentei l , es m a c h t sich eine gewisse H e m m u n g ge l t end : die Blt i ten werden an den Gibbere l l inpf lanzen zwar e twa z u m gleichen Z e i t p u n k t s i ch tba r wie an den d u t c h K u r z t a g indu- z ier ten Kont ro l len , aber es k o m m e n n u r re la t iv wenige Bl i i ten zur E n t w i c k l u n g , n n d n a c h einiger Zeit f inde t ein Riick- sch lag zur v e g e t a t i v e n P h a s e s ta r t . A u c h das L ~ i ngenwachs tum der In te rnod ien , das bet n i ch tb l f i henden K a l a n c h o e p f l a n z e n wesen t l i ch s tbxker i s t als bet b l i ihenden, i s t bet den Gibberel- l inp f l anzen s t a r k gef6rdert , u n d die B l a t t s u k k n l e n z , die bet b l i ihenden P f l anzen besonders hoeh ist, i s t bet den Gibberel l in- E x e m p l a r e n gering. Bet K a l a n c h o e werden die v e g e t a t i v e n Merkma le also du rch Gibberel l in gef6rdert , die r e p r o d u k t i v e n aber g e h e m m t .

Aus dieser g~gzl ich a n d e r s a r t i g e n IReaktion der K u r z t a g - p f lanze K a l a n e h o e gegenf iber u n s e r e n ande ren Versuchsob jek - t e n erg ib t sich, dab das Gibberel l in keinesfal ls der allein a u s s c h l a g g e b e n d e F a k t o r fiir die B l i i t enb i ldung sein k a n n Evgl. ~ ) ] .

M i t U n t e r s t f i t z u n g der D e u t s c h e n F o r s c h u n g s g e m e i n - schaf t .

Gdttingen, P/lanzenphysiologisches Institut der Universit~l t~. HARDXR u n d R. B /3~sow

Eingegangen am 26. November 1956

t BONsow, R., u. R. HARDER: Naturwiss. a) 43, 479 (1956); b) 43, 527 (1956).

fiibberellin and Flower Formation

As r epo r t ed previously1) , gibberel l in induces f lower for- m a t i o n in nonco ld - t r e a t ed b iennia l tfyoscyamus niger, t h u s enab l ing th i s p l a n t to b y p a s s i ts n o r m a l cold r e q u i r e m e n t . T h e e x p e r i m e n t s h a v e meanwh i l e been e x t e n d e d to o the r cold- r equ i r ing p l an t s a n d to a n u m b e r of long- a n d s h o r t - d a y p l an t s , all of t h e m m a i n t a i n e d u n d e r n o n i n d u c t i v e condi t ions . Th i s no t e is to s u m m a r i z e t h e resul ts . A fuller a c c o u n t will be pub - l ished elsewhere.

Cold-requiring plants. Some, b u t n o t all p l a n t s of a v a r i e t y of ca r ro t (Ear ly F r ench Forcing) f o r m e d f lowers a f te r 11/2 to 21/o m o n t h s of da i ly app l ica t ions of t0 [zg or more of g ibberel - lina). I n a n o t h e r v a r i e t y (DANvER'S Ha l f Long), a bo l t ing response on ly ha s been obse rved so far, a n d t h e s a m e is t rue of pa r s l ey (Hamburg ) and t u r n i p s (Purple T op Globe). The t r e a t m e n t is be ing con t inued . In w in te r rye ( P E T K U S ) s t e m e longa t ion was g rea t ly p romo t ed , b u t no effect on f lower

in i t i a t ion was to be de tec ted (same leaf n u m b e r to f i rs t f lower a n d same developmental s t age of the terminal shoot meristem in treated plants and controls)3).

Long-day plants. A rapid flowering response, equalling the effect of long-day exposure, was obtained in Samolus parviflorus with daily applications of 2 ~g or 5 ~g of gibberellin. Flower buds became visible after about one month, open flowers appeared one week later (5 ~zg treatment). A similar r e sponse seems to occur in Crepis tectorum (the e x p e r i m e n t is still in progress) . I n Silent armeria and in a n n u a l ttyoscyamus niger t h e response was m u c h slower a n d so far par t i a l (only p a r t of t he t r e a t ed p l an t s responding , s imi lar to t he Ea r ly F r ench Forc ing carrot) . The m i n i m u m effect ive dose was 10 txg daily, a n d t h e f irst f lower b u d s b e c a m e visible a f te r a b o u t 7 to 8 weeks in Silene a n d af ter f0 weeks in H y o s c y a m u s . In H y o s c y a m u s , t he buds failed to develop fu r t he r ; however , c o n t i n u e d d e v e l o p m e n t m a y be a m a t t e r of p rope r dosage a n d per iods of t r e a t m e n t . I n P e t k u s sp r ing rye, the resu l t s were ident ica l wi th those in nonve rna l i zed win te r rye.

Short day plants. A four week long appl ica t ion of 1 to /0 b~g gibberel l in da i ly g rea t ly e n h a n c e d s t e m e longa t ion in cocklebur (Xanthium saceharatum), b u t t he p l an t s r e m a i n e d s t r i c t ly vege ta t ive . An e x p e r i m e n t wi th s o y b e a n (Biloxi), still in progress , ind ica tes a s imilar behavior .

Comments. I t is ev iden t t h a t gibberel l in is capab le of i n d u c i n g flower fo rma t ion in a considerable n u m b e r of p l a n t s a n d u n d e r s t r i c t n o n i n d u c t i v e condi t ions . However , th i s effect of gibberel l in does no t seem to be general . P r o m o t i o n of f lowering was n o t obse rved in rye (winter a n d spring) nor in s h o r t - d a y p l a n t s kep t in l ong-day condi t ions , a l t h o u g h t h e v e g e t a t i v e effects of gibberell in appl ica t ion were ve ry pro- nounced . W h e r e flower fo rma t ion did occur in b iennia l an d long-day p lan t s , t he response was var iable : r ap id and com- plete in some species, slow a n d poss ib ly pa r t i a l in o thers . This var iab i l i ty c a n n o t be a t t r i b u t e d to differences in photoper iod ic r equ i r emen t , s ince t h e sens i t i v i ty of Samolus and Crepis to l ong -day induc t ion is no t g rea te r t h a n t h a t of H y o s c y a m u s an d Silene. A t present , to a ccoun t for t he var iab le effect of gib- berel l in on f lower f o r m a t i o n two exp l ana t i ons appear m o s t p robab le : 1. Gibberell in, or a s u b s t a n c e of s imi lar physiological ac t iv i ty , is n o t t he sole r egu la to r of f lower fo rmat ion , b u t pa r t i c ipa t e s in th i s process in con junc t ion wi th o the r factors. Thus , t h e fac tors neces sa ry for t he comple te func t ion ing of gibberel l in in f lower fo rma t ion m a y be l imi t ing in some p lan t s , b u t no t in o thers . 2. The gibberel l ins of d i f ferent p l an t s are n o t ident ical a n d no t comple t e ly in te rchangeab le , so t h a t a g iven gibberel l in will n o t be equa l ly effective in all p lan ts , even if gibberel l in is t h e fac tor l imi t ing f lower f o r m a t i o n in t he se p lan t s .

W h e t h e r t h e ac t ion of gibberell in in flower fo rma t io n is d i rec t or indirect , r ema ins to be seen. If t he p l an t s which fo rm flowers af ter gibberel l in t r e a t m e n t are i nduced in t h e n a t u r a l w a y (by a cold t r e a t m e n t or b y long days), bo l t ing a n d flower in i t i a t ion occur a l m o s t s imu l t aneous ly . I n con t ras t , gibberel l in t r e a t m e n t causes s t e m e longat ion to occur first, while f lower in i t i a t ion m a y be more or less delayed. Therefore i t s eems possible t h a t t h e p r i m a r y effect of gibberel l in is ini t ia- t ion of s t e m e longat ion , a n d t h a t f u r t he r processes are se t in m o t i o n in t h e e longa t ing s t e m which in t u r n lead to f lower fo rma t ion . I n th i s case, f lower f o r m a t i o n would be a secon- d a r y effect of gibberell in.

Th i s work was s u p p o r t e d in p a r t b y a resea rch g r a n t (RG-3939) f rom t h e N a t i o n a l I n s t i t u t e s of Hea l th , U.S. Pub l i c H e a l t h Service.

Department of Botany, University o/ California, Los Ange- les 2g, Cal. ANTON LANO

Eingegangen am t6. November t956

t) LANE A.: Naturwiss. 43, 284 (t956). 3) Par t of the experiments was done with a mixture of gibberel-

lin A and gibberellie acid, kindly supplied by Dr. F .H . STODOLA, U. S. D. A . Northern Utilization Research Branch, Peoria, Ill., par t with a preparation containing gibberellic acid only, obtained through the courteousy of Dr. CURT LEBEN, Lilly Research Labora- tories, Indianapolis, Ind.

8) Rye was treated either by applying a drop of gibberellin solu- tion to the base of the blade of the youngest fully expanded leaf, or by immersing a leaf blade, with the tip cut, into the solution. The other plants were treated by applying the solution to the apical region of the shoot. Most experiments were carried out by Mrs. PATRJCIA B. FINN.

Herausgeber und verantwortlicher Redakteur: Professor Dr, Ernst Lamla. G6tfingen. - - Veraixtwortlich ffir den Anzeigenteih Gfinter HoItz, Berlin W 35, Reichpietschufer 20. - - Springer-Verlag, Berlin-Gbttingen-Heidelberg. - - Drnck der Universitiitsdrnckerei H. Sttixtz AG., Wfirzburg. - - Printed in Germany