Volume 136, number 1 FEBS LETTERS December 1981
A PROBABLE LINKING SEQUENCE BETWEEN TWO TRANSMEMBRANE COMPONENTS
OF BACTERIORHODOPSIN
Nandini V. KATRE and Robert M. STROUD
Department of Biochemistry and Biophysics, University of California, School of Medicine, San Francisco, 94143, USA
Received 19 October 1981
1. Introduction
Purple membrane from Halobacterium halobium
contains predominantly one protein, bacteriorhodop-
sin, which acts as a light.driven proton pump [1], and
which consists of 7 transmembrane rods of electron
scattering density [2]. X-Ray diffraction analysis
[3,4] suggests that the rods correspond predominantly
to a-helical regions, oriented roughly perpendicular to
the membrane plane. However, which parts of the
sequence are contained in the rods within the bilayer
and which correspond to the linking regions between
helices is not known. Enzymatic cleavage [5,6] or
chemical modification of the protein, using reagents
which preferentially position either into the aqueous,
or the lipid phase identify linking or transmembrane
regions.
We describe specific labelling oftyrosines accessible
to lactoperoxidase iodination of bacteriorhodopsin
from the aqueous phase. The site of modification is
located by separation of peptides using Sephadex
LH-60 chromatography and amino acid analysis.
2. Methods
2.1. Isolation of purple membrane
Halobacterium halobium strain RI was grown and
purple membrane was isolated as in [7].
2.2. Iodination of tyrosines in purple membrane
The tyrosine residues of bacteriorhodopsin were
modified using sodium [12sI]iodide and the lactoper-
oxidase-glucose oxidase enzyme system [8] immobi-
lized on hydrophilic polyacrylamide beads (Enzymo-
beads), obtained from BioRad. Purple membrane (2
mg) was suspended in 1 ml 0.2 M phosphate buffer
(pH 7.2). The Enzymobead reagent (1 mg) dissolved
in 0.5 ml distilled water was added to the membrane
suspension. A 10-fold excess of NalZSI over purple
membrane was then added and the reaction started
by addition of 0.25 ml 2% glucose. After 15 min at
room temperature, the enzymobeads were removed
by centrifugation 1000 × g for 5 min. The purple
membrane was washed several times with phosphate
buffer, and concentration assayed using extinction
coefficient esT0 = 63 × 10 a cm -1 for the light-adapted
form [9]. The same procedure was used to iodinate
20 mg protein, using non-radioactive Nal.
2.3. Identtfication of labeled residues
The radioactive and non-radioactive iodinated
samples were mixed and initial cleavage of labeled
protein was performed by treatment of apomembrane
with a-chymotrypsin (Sigma Chemical Co.) at a ratio
of 1.5 X 10 -2 mg/mg protein, for 5 h, as described
[5 ]. Apomembrane was generated by bleaching in 1 M
hydroxylamine hydrochloride, pH 7.0 at 37°C for
2-3 hours [10], and collected by centrifugation at
25 000 × g. The two chymotryptic fragments gener-
ated were separated by Sephadex LH-60 chromatog-
raphy (column size 2.0 cm diam. × 60 cm length),
using 96% formic acid and ethanol (2.5:7) as elution
buffer and 2 ml fractions were collected. The larger
of the 2 chains (residues 72-248), named (CTI) and
the smaller chain (CTII residues 1-71) were separated.
CTI was lyophilized, dissolved in 70% HCOOH and
treated with cyanogen bromide (CNBr) in 70%
HCOOH. A 100-fold molar excess of CNBr over
methionine residues was used. The reaction mixture
was incubated at 37°C for 24 h, after which it was
diluted 10-times with distilled water and lyophilized.
Published by Elsevier/North-Holland Biomedical Press
170 00145793/81/0000-0000/$02.75 © 1981 Federation of European BiochemicalSocieties