Euphytica

34 (1985)

785-791

GENETIC

ANALYSIS OF FLOWER AND LATERAL SHOOT PRODUCTTON IN GERBERA

JAN DE JONG and (IVT). FRIDA P.O. Box GARRETSEN 16.6700 AA Wageningen. the Netherlands

Institute

for Horticultural

Plant

Breeding

Gerhrruirm~c,sonii,

gcrhera,

cut flower

yield.

lateral

shoots.

inheritance.

selection

An incomplete dlallel cross with selfs and reciprocals was made Very signilicant dlffcrcnces occurred between GCAs of the number of lateral shoots. The selfs were mainly rcsponsihle for A positive genetic correlation occurs between the number of and total flower production. The phenotypic performance of related to their breeding value (measured on seedlings). It is differences hetwccn cuttings and seedlings. INTRODUCTION

with twelve cultivars of Gwhwu jmzew~rzii. parents for cut llowcr yield. earliness and the signilicant SCAs. lateral shoots at anthesis of the first flower parents (measured on cuttings) was poorly suggested that this is due to physiological

The very short leafy lateral shoots of the Gerbera form sympodia. Generally on each shoot two flowers develop, one from the apical bud and one from the uppermost leaf axil (SHOUB & GLJILI, 1970). An important factor for the yielding capacity of Gerbera cultivars is therefore the number of lateral shoots. This number may be counted early, at the time the first flower buds appear on a seedling (LEFFRING, 1981) or at a later time (HORN et al., 1974). In both cases a positive significant phenotypic correlation has been reported between cut flower yield and number of lateral shoots. Genetic studies on Gerbera report only on cut flower yield, for which a large genetic variation has been observed (DE LEO & OTTAVIANO, 1978; HARDING et al., 198 1; SCHIVA, 1976; WRICKE & HORN, 1971). No information is available on the inheritance of lateral shoot number and only recently a first study on the genetics of the vase life of cut flowers was published (HARDING et al., 19X1). The present experiment was designed to obtain information on the genetics of cut flower yield, days to first flower, lateral shoot number, number of flowers on main axis and vase life of cut flowers. The first paper deals with the inheritance of the first four characters, the second paper will treat of the inheritance of vase life duration.
MATERIALSANDMETHODS

Twelve clones of Gerhera,jumesonii H. BOLUS ex HOOK. differing in keeping quality and yielding capacity were selected as parents for an incomplete diallel cross (almost
785

.I. Uh JONG

ANU

F. G. GAKKETSFIU

balanced) with selfs and with reciprocals. The total number of crosses was 87, including 10 selfs and 27 reciprocals. The parents were obtained from the IVT Ph~tophthova cryptog~a resistance programme (SPARNAAIJ et al., 1975) and from two Dutch Gerbera breeders. Seeds were sown in October 1977 and the seedlings were planted in the greenhouse on raised ground beds at a density of six plants per m2 in January 1978 together with rooted cuttings of the 12 parents. As a consequence, 99 objects were included in the experiment, which was laid out in twelve randomized blocks and one plant per plot. In June 1978 a section of the greenhouse was flooded. which reduced the experiment to six replications from this month onwards. At anthesis of the first flower on each plant the following data were recorded: I) number of days from January 1 to first flower (earliness); 2) number of flower buds on the main axis; 3) number of lateral shoots. Until April 1979 the monthly yield of cut flowers was determined. We calculated the following yield characters: 4) early yield: number of cut flowers per plant for the period from March 1, 1978 to June 1, 1978; 5) winter yield: number of cut flowers from November 1, I978 to April 1, 1979; 6) total yield: number of cut flowers from March 1, 1978 to April 1, 1979. For characters 1 to 6 analyses of variance were carried out separately on the crosses (self included) and on the parental clones. For the characters 3, 4, 5 and 6 first a square root transformation was performed after adding one to the number per plot. Twelve replications were available for analyses of the characters 1. 2, 3 and 4, but only six for characters 5 and 6. The diallel subdivision of the sums of squares of differences between genotypes was made according to the fixed model (GARRETSEN & KEULS, 1978): yI, = p + A*, + 3Ly + S$ + r: + e, in which yl, = mean of the cross between female i and male j, u = general mean, h*, (h:) = GCA of the ilh (j) parent, s*,, = SCA of the cross between i and jth parent, rf = reciprocal effect and e,j = error term. The error MSs per F, mean were derived from the interaction (crosses x replicates) MSs The GCAs in Table 2 were Bonferroni t-tested against their standard errors to show significant positive or negative deviations from zero.
RESULTS The combining

ability analyses of variance with and without selfs presented in Table 1A show that significant GCA-MSs were found for all characters studied. The SCAMSs appeared to be significant or highly significant when selfs were included for all characters except for number of flower buds on the main axis. Comparison of the analyses with and without the selfs showed that the selfs were mainly responsible for the very significant SCA-MSs, however for early yield and days to first flower highly significant SCA-MSs remained after elimination of the selfs. For days to first flower highly significant differences between reciprocals also occurred. Table 1B shows 786
Euphvticcr34 (1985)

2
with (+) and wlthout (-) selfs for five characters m Gcrbera and for one character with sells only. yield Early yield Winter yield Days to lirst flower Number shoots MS + + oflatcral

Table 1. A) Combining B) Analyses of variance

ability analyses of parent clones.

of variance

3 ,-T 2. ;I 2 MS + + + MS MS Number of flowers on main axis MS

Source of variation

Dimension

Total

2 2

MS

A. GCA SCA Rec. Error 1.13** 0.27 0.27** 0.0x** 0.04 0.01 33 0.02 0.26** o.os** 559** 137** 89** 627** X6** 0.21** 0.04* 0.02 0.26** 0.02 0.31** 0.46** 0.50** 0.22* 0.1 I 0.14

0.04* 0.02 0.01 0.01

(per F, mean)

II 48-38 27 406 860

1.01** 0.85** 0.22 0.21

7 C 1 7

B. Between Ps Error (per plant) 0.61** 0.15 309 1.54** 0.46 1363**

II 50Z 105

4.31** 0.56

0.39** 0.08

0.40** 0.16

z E 5 F c 3

Transformed data. Based on six replications *signilicant at P = 0.05;

only. **significant

at P = 0.01.

J. DE JON<;

AND

F. G. GAKKETSEN

Table 2. Estimates of GCAs from the combining ability and lateral shoot number of I2 cultivars ofGerbera. Parent Code Total yield Early yield

analysts

without

sclfs for yield.

days to lirst flower

Winter

yield

Days to first flower I l.6** -3.5 0.5 -x.3** 5.4* 4.6 3.9 x.3** 6.3** -I 1.x** -7.9** 3.6 133.6 I.6

Number of lateral shoots -0.17** -0.1x** -0.14* 0.03 i).O6 0.08 0.08 0.01 0.07 0.14 0.23** 0.2x** 1.62 I.61 0.04

Sympathy Peter Adele Helios Symphony IVT 63007 Lady Oranje Glorlc IVT70159-I8 Ronald IVT 72130-I IVT72123-I Mean Mean

I 2 3 4 5 6 7 8 9 I0 II 12

-0.40* PO.29 -0.28 PO.24 -0.21 0.00 0.00 0.05 0.06 0.35 0.46** 0.5x** 4.86 22.62 0.13

0.17** 0.04 m~o.05 0.11* 0.17** -0.13** -0.04 a.17** 0.15** 0.27** 0.22** -0.0 I .I92 2.62 0.03

-0. IO 0.26 -0.14 0.34** 0.13 0.07 0.07 0.06 0.00 0.19 0.04 0.34** 2.99 7.94 0.1 I

of F,s (transformed) (untransformed) error (GCA) data. significant

Standard

Transformed (**) *: (very)

different

from

0 (Bonfcrroni

t-test)

that the parent clones differ significantly. Table 2 presents the GCA estimates of the parents obtained when selfs are excluded. The GCA estimates for number of flowers on the main axis have been omitted as none of the GCAs was significantly different from 0. The overall mean of the F,s was 2.0 flower buds per plant. The GCA estimates of the parents are presented in order of increasing GCAs for total yield, as this character is commercially important. Table 3 gives in the same order the deviations of the parental clones from their general mean. From Table 2 it appears that for total yield IVT 72130-I and IVT 72123-l have a significant positive GCA estimate and that Sympathy is a parent with a significant negative one. Early yield or winter yield are affected by those and other parents as well. The GCA estimates of the parents for number of lateral shoots correspond in a remarkable way with those for total yield. This close genetic correlation is demonstrated in Fig. 1. None of the GCA estimates of the remaining characters show the comparable close relation with the GCA estimate of total yield. The relation between total yield and number of lateral shoots was not so strongly marked in the parental clones. Only Ronald combined a high total yield with a high number of lateral shoots. The number of shoots of the other clones did not differ significantly from the general mean. Further, comparison of Tables 2 and 3 shows that the genetic behaviour of the parents does not correspond closely with their phenotypic behaviour. The estimated SCAs have not been given but it is concluded from the consistently negative and significant SCA estimates that the selfs were late and less productive, probably as a consequence of inbreeding depression.

CIJT FLOWER

YIELD

OF GhKHEKA

Fig. I The relation between of cultivars in Table 2.

GCAs

for total

cut flower

yield

and early

branching.

Figures

rcfcr

to code

Significant SCA estimates were not only restricted to the selfs but also extended to crosses for the related characters days to first flower and early yield. The crosses Sympathy x IVT 63007 and Peter x Symphony showed significant SCA estimates of 30 and 21 days for days to first flower and of -0.5 and -0.4 for early yield. This means that these crosses were later and had a lower early yield than was expected on the basis of their GCAs. The number of days from January 1 to first flower of the cross Sympathy x IVT 63007 (Y,.~)is thus composed of the following components: Y,.~ = p + h: + A*, + sT6 + &, = ll3,6 + II,6 + 4.6 + 30.3 + 0 = 160.1 Regarding winter yield, the SCA estimate of the cross Oranje Glorie x IVT 70159-18 was significantly negative ((1.3). Reciprocal differences for days to first flower related to IVT 72 123-l. which when used as a male in the crosses with Sympathy and Adele is significantly later than when used as a female, resp. 27 and 22 days.
DISCUSSION

The commercial life span of a Gerbera plant in the Netherlands is up to 1.5 year. Selection for total yield however is possible in an early stage as thischaracter is correlated with number of lateral shoots. LEFFRINC (198 1) and WRICKE & HORN (197 1) arrived at this conclusion on the basis of phenotypic correlations. We conclude from the correspondence between the GCA estimates for flower production and those for lateral EuphJ~tica 34
f 1985)

789

J. I)F JONG

AYD

F. G. GARRETSEN

Table mean. Parent

3. The

parental

clone

means

per plant

for yield,

earliness

and branching

as deviations

from

their

Code

Total

yield

Early

yield

Winter

yield

Days to lirst flower -6.0 -7.3 -I I.9 1.2 -5.7 27.0** -8.6 7.0 3.1 -7.6 6.0 5.7 100.6 5.1

Number of lateral shoots 0.15 -0.16 -0.12 -0.06 0.01 -0. I2 -0.06 -0.13 0.38** 0.23 0.16 I.19 0.42

Sympathy Peter Adele Hclios Symphony IVT 63007 Lady Oranje Gloric IVT70159-IX Ronald IVT72130-I IVT72123-I Mean Mean Standard (transformed) (untransformed) error

I 2 3 4 5 6 7 X 9 IO II I2

-0.48 0.44 -0.19 -0.27 -0.89 -1.15** 0.39 -I .22** 0.14 I .33** 0.x0 I .09 4.X2 22.23 0.30

0.07 0.13 0.12 0.09 -0.40** ~0.42** 0.04 -0.09 -0.14 0.33* 0.19 0.04 2.10 3.41 0.1 I

0.26 -0.06 -0.18 -0.48 -0.05 -0.54 0.43 0.83 -0.24 0.53 0.22 0.98* 2.90 7.4 I 0.28

(per plant) data. significant

Transformed (**) *: (very)

different

from

0 (Bonferroni

t-test)

shoot number that there is also a genetic correlation. Lateral shoots may sprout from nodes anywhere along the main stem. The closer to the apex a lateral shoot originates, the fewer leaves it forms below the flower and the earlier it may flower (LEFFRING, 1981). Measurements in our experiment, not reported here, failed to show significant differences between the progenies in the node number from which the second lateral shoot originated ~ the first always comes from node two -; thus in our material this character was unfit for selection. The correspondence between the breeding value of the parents with their phenotypic performance as a clone was not very close, although the three best branching clones were also the parents that transmitted good branching to their progenies. The poor correspondence may be attributed to physiological differences between seedlings, on which the breeding value is based, and cuttings from which data the phenotypic performances are derived. LEFFRING (1981) reports that the seedling main growing point produces 7 to 24 leaves before the flower bud is initiated, against 2-9 for the lateral growing points. Cuttings which are rooted lateral shoots taken from adult plants, are more likely to respond as lateral shoots than as the main axis of a seedling. The corresponding smaller leaf initiating capacity most likely affects earliness and branching capacity. In our experiment the cuttings flowered ten days earlier than seedlings possibly because the flower buds were partly initiated while on the mother plant. At anthesis of the first flower the cuttings had produced only 0.42 lateral shoots per plant versus 1.61 for the seedlings. SEECHERS (pers. corn.), in his Gerbera breeding material, has often observed that seedlings have more lateral shoots than cuttings at anthesis of the first flower. Cuttings ultimately produce as many laterals but branching commences somewhat later. It seems therefore wise to count the number of lateral shoots 790
Euph~lica 34 (1985)

on cuttings not at, but after anthesis of the first flower. Only then may the inherent branching and yield capacity of a cutting be properly estimated.
REFERENCES Dr: LEO. V. & E. OTTAVIANO. 1978. Genctical analysis of morphological traits in Gerhercr jummmii. Clonal and diallel families. Eucarpia Meeting on Carnation and Gerbera. Alassio: 179-191. GAKKETSEN. FKIUA & M. KFIJLS. 1978. A general method for the analysis of gcnctic variation in complete and incomplctc diallels and North Carolina II (NC II) designs. Part II. Proccdurcs and general formulas for the lixed model. Euphytica 27: 49-68. HAKUINC;, JAMES, THOMAS G. BYRNE & R~~EKT L. NELSON, 1981. Estimation of heritability and response to selection for cut-flower yield in Gerbcra. Euphytica 30: 3 13-322. HAKI)IN~;, JAMES. THOMAS G. BYKNE & ROB~K~ L. NELSON. 1981. Heritability of cut-llowcr vast longcvlty in Gerbera. Euphytica 30: 653-657. HORN. W.. G. WKICI<E & W. E. W~BEK. 1974. Erbliche und umweltabhangige Merkmalausprigung bei Grrhcrtr jum~sonii. Gartenbauwissenschaft 39: 2X9-300. LEFIKIN~;, L., I98 I. De bloemproduktie van Gerbera. Thesis Agricultural University. Wageningen, pp.
X6.
SCHWA.

T., 1976. Gcrbcra breeding. Preliminary evaluations of genotypes for improved populations of production. Acta Horticulturae 63: 177-185. SHOLIH. J. & A. C~IJILI. 1970. A new system for vegetative propagation of Gcrbcra. Proc. XVIII Int. Hort. Congr. I: 232. SPAKNAAIJ, L. D., FKIIIA GAKKE~S~N & WIES BEKKEK, 1975. Additive inheritance of resistance to Pl?~,/o$~horcr cr?pfo~~m PETHYBKIU(;E and LAFFEKTY in Gwherrr jumwnii Bo~.us. Euphytica 24: 55 l-556. WRI(.KE, G. & W. HORN. 1971. Gcnetische und ziichterische Untcrsuchungen bei Grrhcrrr. Eucarpia Meeting on Ornamentals. Wagcningen: X5-98.

791