Worm Breeder's Gazette 5(2): 16

These abstracts should not be cited in bibliographies. Material contained herein should be treated as personal communication and should be cited as such only with the consent of the author.

Triple Helical Collagens in the Adult Cuticle of C. elegans

M. Kusch, Z. Kahn

Studies on vertebrate collagens have revealed that pepsin, at low 
temperature and pH- specifically digests non-helical regions (usually 
at NH2 and COOH termini) of native collagen, but does not digest 
triple helical regions.  Since C.  elegans cuticles contain 70% 
collagen (by weight), we decided to investigate the effects of limited 
pepsin digestion.
Our procedure for isolating cuticle proteins involves 3 steps: first,
sonication of whole animals in a low osmotic strength buffer; second, 
boiling these crude cuticles in 1% SDS for 2 min to remove all non-
cuticle material; and third, solubilization of cuticle proteins with  -
mercaptoethanol in the presence or absence of detergent.
In separate experiments cuticle proteins were exposed to pepsin at 
each of these three steps:  After sonication but before incubation 
with SDS; after incubation with SDS followed by extensive washing to 
remove detergent; after extraction with  -mercaptoethanol in a neutral 
salt solution.
With either sonicated or SDS-cleaned cuticles, cuticle proteins were 
not solubilized by pepsin but were solubilized after subsequent 
reduction of S-S bonds with  -mercaptoethanol.  SDS-PAGE of the 
solubilized proteins separated them into 8 bands (MW range: 40K-125K). 
These bands were identical in sonicated and in SDS-cleaned cuticles 
suggesting that SDS does not irreversibly denature the proteins.  In 
controls without pepsin the usual cuticle protein pattern was 
observed: 8 bands with molecular weights between 54K and 212K and 
large numbers of higher molecular weight bands which account for 20% 
of the soluble proteins.  No high molecular weight material was 
present in gels of pepsin-treated proteins suggesting that the high 
molecular weight is composed of the low molecular weight (54K-212K) 
species crosslinked within the non-helical regions.  Pepsin digestion 
of these regions presumably converts them into the eight 40K-125K 
pepsin resistant proteins.  In addition, if pepsin actually degrades 
all nonhelical material, these experiments indicate that S-S bonds 
within helical regions are responsible, at least in part, for 
crosslinking of collagens within the cuticle since  -mercaptoethanol 
is still required to solubilize the pepsin resistant material.
In a third type of experiment, SDS was washed out of cleaned 
cuticles and proteins were extracted with  -mercaptoethanol into a 
neutral salt solution and then exposed to pepsin in order to see if we 
could extract native helices that are pepsin resistant when not 
attached to the cuticle.  Resistance to pepsin degradation was used as 
a criterion for helix stability.  SDS-PAGE of these digestion products 
gave 7 bands (MW 39K-110K).  The molecular weights and relative 
amounts of these proteins appeared similar but not identical to those 
obtained when in situ cuticle proteins were exposed to pepsin.  In all 
three types of experiments 2 very prominent major bands were obtained (
MW 67K and 81K).  Non-pepsin-treated control cuticle extracts also 
contain 2 very prominent bands (MW 91K and 106K).  It is tempting to 
speculate that pepsin simply degrades the non-helical termini of these 
proteins to create the two lower molecular weight bands.  If this is 
true, then approximately 25% of each of these collagen chains is not 
in a triple helical conformation.  This figure is in good agreement 
with estimates of the maximal possible amount of triple helix (78%) 
from determinations of the glycine content of cuticle collagens (Cox, 
Kusch and Edgar, manuscript submitted).  The pepsin resistance of 
solubilized cuticle proteins demonstrates that native collagen helices 
can be extracted from cuticles by reduction of S-S bonds.  Although 
interchain S-S bonding occurs within helical regions, presumably this 
crosslinking is not necessary for maintenance of native helices.
These experiments indicate that C.  elegans cuticle collagen like 
vertebrate collagens, is in a triple helical conformation.  In C.  
elegans, however, these helices are cross-linked by S-S bonds.  
Molecular weights of the pepsin-resistant polypeptides suggest that a 
single polypeptide is not folded upon itself to form a triple helix, 
as has been suggested for Ascaris cuticle collagen