Activity of anthocyanin biosynthetic enzymes and CHS in anthocyanin deficient mutants in tomato.

Jorgensen, R.A., and Dooner, H.K.

The present study was undertaken with the hope of identifying the genetic loci encoding the anthocyanin biosynthetic enzymes UDPglucose 3-0-glucosyltransferase (UFGT) and chalcone synthase (CHS) . Mutations in the genes encoding these enzymes in maize, petunia, and other species result in a recessive anthocyaninless phenotype. Since these genes have been molecularly cloned from maize and petunia, it is possible to introduce them to tomato by transformation and test for complementation of the homologous tomato genes. As a prerequisite to performing this test, we set out to identify mutants of these enzymes in tomato by enzyme assay. There are eight known genetic loci in tomato with alleles conditioning a "completely anthocyaninless" phenotype (a, aa, ae, af, afr, ah, aw, and bls; obtained from C. Rick); afr and bls have pleitropic effects on- growth, the other six "completely anthocyaninless" mutants have not been reported to possess any other altered characteristics besides loss of anthocyanin pigmentation, and so are clearly candidates for anthocyanin biosynthetic enzyme genes. We assayed UFGT and CHS activity in leaves from seedlings that had been grown for 13 days under high intensity fluorescent lighting (16h. days, 8h nights) at 16C, conditions that induce high levels of anthocyanin production. UFGT was assayed in extracts prepared from 50 mg leaf tissue (not fully expanded) homogenized in 100 ul cold 35mMtris8.2 and diluted to 200 ul. 25 ul of each extract was assayed for UFGT as described by Dooner (1981). Reactions were 50ul and were extracted with 300 ul ethyl acetate. 200 ul of the ethyl acetate phase was counted in a scintillation counter (see Table 1). Controls were run without quercetin. CHS was assayed in a similar homogenate except for the use of 0.1 M KPO4, pH 7.6, 1.42 mM B-mercaptoethanol and the addition of BSA to 5 mg/ml immediately after homogenation. 40 ul homogenate was assayed as described in Dooner (1983) in 50 ul. The reaction was stopped with 20 ul methanol and extracted with 300 ul ethyl acetate. Controls were run without p-coumaroyl CoA. 150 ul ethyl acetate phase was counted (Table 1) and 100 ul was spotted on Whatman #1 paper and chromatographed in 30% HOAC to confirm the identity of the naringenin reaction product.

     Table 1.  Activities (in cpm) of UFGT and CHS.
____________________________________________________________
Genotype  Accession            UFGT  Control   CHS  Control
_______   _________            ____  _______   ___  _______
Control   "Red Cherry" Herbst  3105       47  1282       77
a/a       LA784                6518       10   876      136
aa/aa     LA1194               3325       30   183       98
aa/aa     LA1525               5195       25   289      134
aa/aa     LA1700               5655       40  4495      215
ae/ae     LA1665               6810       40   694      115
af/af     LA1444               2078       10   998      117
afr/afr   LA1784               7458       38  1414      119
ah/ah     LA1164               2490       33  1840      133
aw/aw     LA271                3655       35  1965      191
bls/bls   LA1004               1295       33   430      108
We find no evidence that any of these anthocyanin deficient mutants lacks either UFGT or HS activity. Although some genotypes possess lower levels of enzyme activity than the "Red Cherry" control, these differences are not sufficient to account for the lack of anthocyanin in these plants. They probably result from the various genetic backgrounds these mutants are in. For example, 2 of the 3 aa/aa accessions showed low CHS activity, but the third one was even higher than the control. We also conclude that none of these genes can be regulatory loci like R and B in maize since none results in null activities for both enzymes.

Literature Cited:

Dooner, H.K. 1981. CSHSQB. 45:457-462.

Dooner, H.K. 1983. Mol. Gen. Genet. 189:136-141.