Acta Zoologica (Stockholm) 82: 261–274 (October 2001)
© 2001 The Royal Swedish Academy of Sciences
Abstract
Cima, F., Perin, A., Burighel, P. and Ballarin, L. 2001. Morpho-functional
characterization of haemocytes of the compound ascidian Botrylloides leachi
(Tunicata, Acidiacea). — Acta Zoologica (Stockholm) 82: 261–274
A morpho-functional study of the colonial ascidian Botrylloides leachi haemocytes
was carried out to propose their classification, relationships and specializations.
This characterization was obtained by (i) investigations of both living and aldehyde-
fixed cells by light and electron microscopy; (ii) cytochemical and cytoenzymatic
assays; (iii) lectin-affinity assays; (iv) phagocytosis and haemagglutination assays;
and (v) anti-CD34 immunocytochemical assay for vertebrate haematopoietic
stem cells. Results indicate that the haemoblast is a circulating stem cell and there
are at least five haemocyte differentiation pathways, the last two of which have
never been described in botryllids: (i) phagocytic line (hyaline amoebocytes and
macrophage-like cells) share ultrastructural features, the same hydrolytic enzymes
and WGA lectin binding, and are involved in yeast phagocytosis and erythrocyte
rosette formation; (ii) cytotoxic line (granular amoebocytes and morula cells)
with vacuoles containing oxidative enzymes and polyphenolic substrates;
(iii) vacuolated cell line (pigment cells and nephrocytes) involved in catabolite
storage; (iv) compartment cell line (compartment amoebocytes and compartment
cells) able to agglutinate erythrocytes and characterized by vacuoles with a
moderately electron-dense content, positive to arylsulphatase activity and binding
DBA, UEA-I, HPA lectins; and (v) granular cell line includes trophic cells,
able to infiltrate the gut epithelium, showing a cytoplasm filled of PAS-positive
vacuoles with arylsulphatase, chloroacetylesterase and β-glucuronidase activities.
Francesca Cima, Department of Biology, University of Padova, Via Ugo Bassi
58/B, I-35121 Padova, Italy. E-mail: ascilab@civ.bio.unipd.it
Blackwell Science, Ltd
Morpho-functional characterization of haemocytes
of the compound ascidian Botrylloides leachi
(Tunicata, Ascidiacea)
Francesca Cima, Andrea Perin, Paolo Burighel and Loriano Ballarin
Department of Biology, University of
Padova, Via Ugo Bassi 58/B, I-35121
Padova, Italy
Keywords:
Botrylloides leachi, haemocytes, ascidians,
ultrastructure, enzymatic activity
Accepted for publication:
30 November 2000
Introduction
In the last few decades, interest in invertebrate immune
responses has led to a great increase in the scientific production
dealing with haemocytes and coelomic cells, their classification,
mutual relationships and biological roles (see Ratcliffe and
Rowley 1981; for reviews).
As regards ascidians, although there is abundant literature
describing the morphology of both living and fixed haemocytes
(Pérès 1943; Endean 1955; Sabbadin 1955; Andrew 1961; Smith
1970a,b; Schlumpberger et al. 1984; Sawada et al. 1991, 1993;
Dan-Sohkawa et al. 1995) and their ultrastructure (Overton
1966; Milanesi and Burighel 1978; Scippa et al. 1982; Burighel
et al. 1983; Zhang et al. 1992; Azumi et al. 1993; Sawada et al.
1993; Sugino et al. 1993), many doubts and uncertainties still
persist on both their classification and differentiation pathways,
mainly due to the adoption of different classification criteria
for each species studied. Unifying classification schemes
have been proposed (Goodbody 1974; Wright 1981; Rowley
et al. 1984; De Leo 1992; Burighel and Cloney; 1997). Many
biological functions, other than defence reactions, have been
ascribed to ascidian haemocytes, such as excretion, nutrition,
budding, germ cell formation and tunic synthesis (Goodbody
1974; Wright 1981; Burighel and Cloney 1997).
In botryllid ascidians, studies of the morphology and
ultrastructure of haemocytes mainly focus on species of the