Fibrinolysis(I 996) 10( I ), 49-63
© Pearson Professional Ltd 1996
The role of carbohydrate in
the activation of plasminogen
2 glycoforms by streptokinase
S. R. Pirie-Shepherd, R. L. Serrano, N. L. Andon,
M. Gonzalez-Gronow, S. V. Pizzo
Department of Pathology, Duke University Medical Center, Box 3712, Durham, North Carolina, 27710, USA.
Summary Plasminogen 2 has six giycoforms that differ in their sialic acid content. In this study we have examined the
effect that this differential sialylation has on activation of these glycoforms by streptokinase (SK). We find that increases
in sialic acid content, above a threshold level, result in a decrease in the catalytic efficiency of activation. This decrease
in catalytic efficiency appears to be due to an increase in Km values and a decrease in/(ca t values. This phenomenon is
most apparent in the activation of plasminogen 2¢, the most extensively sialyiated glycoform with 13 mols of sialic
acid/mol protein. Gel filtration demonstrated that SK forms an activator complex with plasminogen 2t), indicating that the
carbohydrate on plasminogen 2 interferes with the stability of the Michaelis complex, ra, ther than the ability of
streptokinase to form activator complexes with plasminogen 2 glycoforms. For comparison we also show data
pertaining to activation of plasminogen 2 giycoforms by urinary plasminogen activator and tissue-type plasminogen
activator.
INTRODUCTION
Plasmin, the central proteinase in fibrinolysis, is generated
from plasminogen, which exists in two main glycosylated
variants. ]-a Plasminogen 1 contains two carbohydrate
chains, one N-linked to Asn-289 and the other a mucin-
type O-linked oligosaccharide at Thr-345, whilst form 2
contains only the carbohydrate chain linked to Thr-345.
Further studies of the glycoforms have demonstrated 6
glycoforms of plasminogen 24 termed tx, 13, 7, 8, e and
with pI values ranging from approx, pH 8.2 to pH 6.6. The
microheterogeneity stems from variation in sialic acid con-
tent of plasminogen, s Recent work from this laboratory4
has demonstrated that the sialic acid content affects the
activation of plasminogen by tissue-type plasminogen acti-
vator (t-PA) bound to a fibrin surface, as a general rule, the
greater the sialic acid content, the lower the catalytic effi-
ciency of activation of the plasminogen glycoforms.
Received: 19 September 1995
Accepted after revision: 11 December 1995
Correspondence to: S. R. Pirie-Shepherd, Duke University Medical Center,
Box 3712, Durham, North Carolina, 27710, USA. Tel: 1-919-684-8986;
Fax: 1-919-684-8689.
Plasminogen can also be activated by urinary-type
plasminogen activator (u-PA) and streptokinase (SK). The
activity of t-PA is largely confined to the vasculature
owing to its marked affinity for fibrin, which facilitates its
physiologic activity on clot lysis. Monocyte cells provide
binding sites for plasminogen and u-PAil The u-PA recep-
tor (u-PAR) is a GPI-anchored protein that serves a dual
function localizing u-PA and mediating the internaliza-
tion of u-PA complexed to its physiological inhibitor plas-
minogen activator inhibitor 1 (PAI-1) (see review ref. 6).
The activation of bound plasminogen is kinetically
enhanced by cell bound u-PA, and the generated plasmin
is also resistant to inhibition by ct2-antiplasmin7 (ct2-AP).
SK is a 414 amino acid, 47kDa extracellular protein
secreted by hemolytic streptococci. 8 SK forms an
equimolar complex with plasminogen (Pg-SK) or plasmin
which acts as an activator of plasminogen. 9 SK has no
inherent proteolytic activity.I° The Pg-SK complex catal-
yses the conversion of free plasminogen to plasmin, a
functional activity that free plasmin does not possess. 9
Recent work has attempted to elucidate the regions of
SK responsible for binding plasminogen. Rodriguez et
al ]~ demonstrated that a 17 kDa fragment of SK, was the
smallest stable plasminogen binding domain to be
49