ISOLAMENTO DE UMA FRAO ENRIQUECIDA EM SINAPTOSOMAS DE CREBRO DA ABELHA APIS MELLIFERA UTILIZANDO GRADIENTE DESCONTNUO DE PERCOLL E IDENTIFICAO DE MIOSINAS V E VI


Original Article 181

FORMATION OF THE SPERMATOZOON DURING AND AFTER 
SPERMIOGENESIS IN Iguana iguana LINNAEUS, 1758 (Reptilia, Iguanidae), 
WITH SPECIAL ATTENTION TO THE MIDDLE PIECE AND AXONEMIC 

COMPLEX 
 

FORMAÇÃO DO ESPERMATOZÓIDE DURANTE E APÓS A ESPERMIOGÊNESE 
EM Iguana iguana LINNAEUS, 1758 (Reptilia, Iguanidae), COM ESPECIAL 
ATENÇÃO PARA A PEÇA INTERMEDIÁRIA E COMPLEXO AXONÊMICO 

 
Adelina FERREIRA1

1. Professora, Doutora, Instituto de Biociências, Departamento de Biologia e Zoologia, Universidade Federal de Mato Grosso, Cuiabá, 
MT, Brasil. adelina@ufmt.br  

 
ABSTRACT: Spermiogenesis involves a series of cellular modifications, culminating in the production of a 

highly specialized cell, the spermatozoon. To reassess the peculiarities of this process, specimens of the lizard Iguana 
iguana were collected in the Pantanal, Mato Grosso do Sul state, Brazil; the testis and ducts were processed according to 
routine methods for light, scanning and transmission electron microscopy. After nuclear condensation and elongation, the 
cytoplasm and organelles flow towards the basal area, below the nucleus. An infolding occurs at the nuclear base, where 
the middle piece is implanted. This area contains a centriole pair, with the proximal one perpendicular and the distal one 
parallel to the nuclear axis. The distal centriole can be functionally compared to the basal corpuscle at the axoneme base. 
The mitochondria, that are round or elongated with transverse cristae, initially organized around the centriole in continuous 
rings; later they alternate with dense bodies. Thus the dense bodies are apparently formed by mitochondrial modification 
with deposition of dense material. In I. iguana the formation of dense bodies is one of the last events of spermiogenesis. 
Marking the transition between middle piece and flagellum is the annulus. This structure blocks the displacement of 
mitochondria from the middle piece during flagellar movement. Halfway down the middle piece, a fibrous sheath is 
formed by the accumulation of an amorphous layer surrounding the axoneme, and this structure is the axonemic complex. 
In the spermatozoa of I. iguana, the flagellar end piece is reduced in diameter, because the fibrous sheath terminates and 
only the typical axonemal microtubule pattern (9+2) is present. A large amount of cytoplasm is observed around 
spermatozoa in the seminiferous tubule lumen, suggesting that these are "immature" spermatozoa. Structural modifications 
occur during their passage through the reproductive tract. These post-testis modifications have not been precisely located 
along the reproductive tract and are probably not essential for spermatozoon mobility. Some features of the male 
reproductive tract are shown. 

 
KEYWORDS: Reproduction. Lizard. Ultrastructure. Seminal receptacle. 

 
 
INTRODUCTION 

 
The modification of testicular germ cells to 

give rise to spermatozoa in lizards has already been 
explored (JAMIESON, HEALY, 1992; HEALY, 
JAMIESON, 1994; JAMIESON, 1995; JAMIESON 
et al., 1996; OLIVER et al., 1996). The reproductive 
tract contributes with the production of secretion 
and the final maturation of spermatozoa in lizards 
(DUFAURE, SAINT GIRONS, 1984; DEPEIGES 
et al., 1985; DUFAURE et al., 1986; HAIDER, 
RAI, 1986; RAVET et al., 1987; COURTY, 1991). 
However, this process has not been detailed for 
Iguana iguana, with an important description of the 
spermatozoon for Vieira et al. (2004). This 
important representative of the Iguanidae family has 
ample distribution in neotropical regions (ZUG, 
1997) and is representative of the Brazilian Pantanal 
fauna. The lizard is large, with the young measuring 

about 15 centimeters while adults reach 2 meters in 
length (FERREIRA et al., 2002a). They present a 
peculiar seasonal reproductive cycle adapted to the 
environmental conditions of the Pantanal region 
(FERREIRA et al., 2002a).  

The spermiogenesis of this species includes 
a few more complicated structures, described in 
Lacertilia, usually known as the Acrosomal complex 
and the Axonemic complex (FERREIRA, 2002b). 
The acrosomal complex, is made up of the 
acrosome, the perforatorium and other associated 
structures, which coat the apical surface of the 
spermatozoon head (FERREIRA et al., 2002b). 
Here, the reproductive process was investigated to 
describe the axonemic complex, including features 
of the final stage of spermiogenesis, as well as the 
reproductive duct’s epithelium. 
 

Biosci. J., Uberlândia, v. 25, n. 2, p. 181-187, Mar./Apr. 2009 

mailto:adelina@ufmt.br


Formation of the…   FERREIRA, A. 182

MATERIAL AND METHODS 
 

Specimens of the lizard Iguana iguana were 
collected in the Pantanal of Mato Grosso do Sul 
(14o22oS and 53o61oW) Brazil. They were 
anesthetized with ethyl ether inhalation, the testis 
and reproductive tract removed, fixed and processed 
according to routine methods for scanning (SEM) 
and transmission electron microscopy (TEM) and 
for light microscopy (LM), stained with 
hematoxylin-eosin and periodic acid Schiff (PAS). 
 
RESULTS AND DISCUSSION 

 
During the reproductive period of Iguana 

iguana, seminiferous tubules contain lots of germ 
cells in different developmental stages and a large 
amount of spermatozoa in the lumen (Figs 1A, 1B).  
During spermiogenesis one of the more complicated 
structures is the axonemic complex, formed by the 
middle piece and flagellum (Fig. 1C). A pair of 
centrioles is found in the contact region of head and 
flagellum (Fig. 1D), the proximal one lying 
perpendicular while the distal one is parallel to the 
longer spermatozoon axis (Figs. 1E, 1H). These 
centrioles are similar to the basal corpuscles of cilia 
in that they organize the microtubule arrangement of 
the axoneme (JAMIESON, HEALY, 1992; 
JAMIESON, 1995; ALBERTS et al., 1994). 
Completing the middle piece a number of 
mitochondria are directed towards this region 
surrounding the centrioles (Fig. 1F) in a circular 
arrangement (Fig. 1E, 1G, 1H, 1J). Between this 
ring of mitochondria and the initial portion of the 
axoneme, a cylindrical layer of peripheral fibers 
appears (Fig 1E, 1J). The last structures to be 
formed in the middle piece are the dense bodies 
between the mitochondria (Figs. 1G, 1H, 1J).  The 
middle piece is limited by a ring constricting the 
cytoplasm known as annulus (Figs. 1E, 1J). The 
middle piece in lizards varies greatly in size 
(JAMIESON, 1995; OLIVER et al., 1996). Also, the 
order of formative events is variable and some 
authors describe dense bodies formed previous to 
the mitochondria alignment around centrioles 
(PHILLIPS, ASA, 1993). The flagellar axoneme has 
the typical 9+2 microtubule arrangement (Fig. 1J, 
1K). Extending along the axoneme is a fibrous 
sheath made up of regularly distributed separate 
rings (Figs. 1E, 1K). The fibrous sheath rings are 
electron dense and amorphous, covered externally 
by the plasma membrane. Elastic properties have 
been attributed to the sheath, which would give it an 
important function in spermatozoon mobility 
(FAWCETT, 1970; SOLEY, 1994; JAMIESON, 

1995, FERREIRA, DOLDER, 2003). In the final 
flagellar region the fibrous sheath progressively 
reduces its thickness so that the final portion of the 
flagellum consists only of the axoneme (Fig. 1L). 
This feature is common to various lizards 
(JAMIESON et al., 1996, TEIXEIRA et al., 1999, 
FERREIRA, DOLDER, 2003). The spermatozoa 
eliminated into the seminiferous tubule lumen still 
present a large amount of cytoplasm around the 
head region but not in the flagellum (Fig. 1I). This 
feature has not been previously reported in the 
literature (FERREIRA et al., 2002a). 

The reproductive tract epithelium contains 
secretory and basal cells. During the reproductive 
period the epithelium thickness varies noticeably 
depending on the size of these cells and the amount 
of secretion produced (Figs. 2A, 2B). Cell secretions 
consist of PAS-positive glycoproteins and 
glycosaminoglycans (Figs. 2D, 2E) contained in 
large vesicles (Figs.s 2C, 2E, 2F). Secretory regions 
carry out a function similar to mammal seminal 
vesicle and prostate (DUFAURE et al., 1986, 
FERREIRA, DOLDER, 2007). This is common in 
other lizards, with variations in the intensity and 
granule volume (DEPEIGES, DUFAURE, 1977; 
DUFAURE, SAINT GIRONS, 1984; DEPEIGES et 
al., 1985; HAIDER, 1985). This secretion is 
responsible for an increase in volume of spermatic 
fluid, permitting mobility and preventing 
spermatozoon dispersal (DUFAURE, SAINT 
GIRONS, 1984; DUFAURE et al., 1986; MESURE 
et al., 1991, FERREIRA, DOLDER, 2003; 
FERREIRA, DOLDER, 2007). It is only in the 
reproductive period that the duct’s epithelium 
reaches its maximum height, as well as maximum 
activity (KOHANE et al., 1980; MESURE et al., 
1991). 

In the epididymis dense bodies in the 
middle piece reach their final shape (Figs. 2G, 2H, 
2I). The final duct region widens to form a glandular 
structure called the seminal receptacle (Figs. 2J, 
2L), which also produces an intense glycoprotein 
secretion (Fig. 2K). This structure is common in 
lizards (CONNER, CREWS, 1980; KUMARI et al., 
1990; SRINIVAS et al., 1995; TOKARZ, 1999; 
FERREIRA, DOLDER, 2007) and snakes (FOX, 
1956; SCHAEFER, ROEDING, 1973; HALPERT 
et al., 1982) and is responsible for the storage and 
final spermatozoon characterization, adding surface 
glycides and probably antigens, essential to 
successful reproduction (CONNER, CREWS, 1980; 
ADAMS, COOPER, 1988; OLSSON et al., 1994; 
FERREIRA, DOLDER, 2003). 

 

Biosci. J., Uberlândia, v. 25, n. 2, p. 181-187, Mar./Apr. 2009 



Formation of the…   FERREIRA, A. 183

 
Figure 1: A. and B.  Seminiferous tubules viewed during the reproductive cycle of Iguana iguana, with the presence of 

developing germ cells and many spermatozoa (asterisks). A. Light microscopy (LM), hematoxylin-eosin 
stained. X600. B. Scanning Electron Microscopy (SEM) X800. C. and D. SEM of spermatozoa. Head region 
(h), middle piece (mp). Arrow indicates the flagellum implantation region. X8000. E-L: Transmission Electron 
Microscopy (TEM) of advanced spermatids and spermatozoa in the lumen of seminiferous tubules. E. Nucleus 
(n), mitochondria (m) forming rings around proximal and distal centrioles; middle piece limited by the annulus 
(arrow), and flagellum formed by the axonemic complex (ac). X14000. F. The arrows indicate the streaming of 
residual cytoplasm towards the tip and mitochondria (m) to the middle piece; ribosomes (r). X 12000. G. Initial 
formation of dense bodies between mitochondria (arrows). Dense fibers of the axonemic complex (ac) appear 
half may down the middle piece . X16000. H. Slightly dense bodies (arrows) around the proximal centriole (pc) 
and distal centriole (dc). X18000. I. Spermatozoa in the seminiferous tubule lumen, with a great amount of 
cytoplasm around the head (asterisks) but none around the flagellum (arrows). X3000 J., K. and L. Transverse 
sections of different regions of the axonemic complex: J. Middle piece: in the center the axoneme (ax), is 
surrounded by the peripheral fibers (pf), annulus (an), mitochondria (m) and dense body (arrow). X48000. K. 
Principal part of the flagellum including axoneme (ax), surrounded by the fibrous sheath and plasma membrane 
(p). X64000. L. Terminal part of flagellum consisting in only the axoneme (ax). X72000. 

Biosci. J., Uberlândia, v. 25, n. 2, p. 181-187, Mar./Apr. 2009 



Formation of the…   FERREIRA, A. 184

 
Figure 2: A. and B. LM of the epididymis of Iguana iguana, hematoxylin-eosin stained. X400. A. At the beginning of 

the reproductive cycle, with reduced secretion (arrows) and absence of spermatozoa in the lumen. B.  In the 
reproductive period, with many spermatozoa (z) in the lumen well and developed secretory cells (arrows). C. 
SEM of the epididymis in the reproductive period. The arrows indicate the abundance of large secretory 
vesicles.  X500. D. Final region of the epididymis, with spermatozoa (z) surrounded by PAS positive 
secretion (s). X200. E. Detail of the epididymal epithelium, the arrow indicates secretion vesicles. X1000. F. 
TEM spermatozoon in the epididymis; arrows indicate the dense bodies, a little more developed. Around the 
spermatozoon are many secretory vesicles (s). X12000. G. H. and I. Transverse sections of the middle piece 
where arrows indicate the dense bodies in stages of differentiation. X40000 J. LM of seminal receptacle, 
showing the glandular aspect, supported by connective tissue septae (arrows), branching out from the capsule 
(c). X100. K. Demonstration of PAS positive secretion (s) and apical region epithelial cells of the seminal 
receptacle (arrows). X400. L. SEM of the seminal receptacle. The arrow indicates the aperture region of the 
cloaca. Internally, the organ forms crypts (cr) where the spermatozoa are stored until copulation. X200. 

 
CONCLUSION 
 
In conclusion, during spermiogenesis in I. 

iguana the basic structures necessary for 
spermatozoon function are formed. The loss of 
excessive cytoplasm as well as the completion of the 
middle piece dense bodies apparently occurs during 

the passage though the reproductive ducts. The duct 
cells and those of the seminal receptacle produce a 
great amount of secretion that contributes to the 
final spermatozoon maturation. Glycoproteins 
secreted by the duct’s epithelial cells are probably 
added to the spermatozoon surface.  
 

Biosci. J., Uberlândia, v. 25, n. 2, p. 181-187, Mar./Apr. 2009 



Formation of the…   FERREIRA, A. 185

ACKNOWLEDGMENTS: We thank the 
IBAMA for granting permission (proc. # 
02014.0077/01-10) to collect a few Pantanal lizard 
specimens, since Brazilian law protects these 

animals. This study was supported by FAPESP 
(proc. # 00/03259-5) and Fapemat (proc. # 
0769/2006). 

 
 

RESUMO: A espermiogênese envolve uma série de modificações celulares, culminando na formação de uma 
célula altamente especializada, o espermatozóide. Para re-avaliarmos as peculiaridades desse processo, espécimes do 
lagarto Iguana iguana foram coletados no Pantanal, estado de Mato Grosso do Sul, Brasil; os testículos e ductos foram 
retirados e processados de acordo com a rotina para microscopia de luz e eletrônica de transmissão e varredura. Após a 
condensação e alongamento do núcleo, o citoplasma e as organelas se deslocam para região basal abaixo do núcleo. Nessa 
região basal do núcleo é implantada a peça intermediária. Essa área contém um par de centríolos, sendo um proximal e 
perpendicular e um distal paralelo ao eixo maior do núcleo. O centríolo distal pode ser comparado funcionalmente ao 
corpúsculo basal na base do axonema. As mitocôndrias são redondas ou alongadas com cristas transversais, organizadas 
em torno dos centríolos em anéis contínuos; esses anéis são alternados por corpos densos. Esses corpos densos são 
aparentemente formados por mitocôndrias modificadas com deposição de material denso. Em I. iguana a formação desses 
corpos densos é um dos últimos eventos da espermiogênese. O que marca a transição entre a peça intermediária e o flagelo 
é o annulus. Esta estrutura obstrui o deslocamento das mitocôndrias da peça intermediária durante o movimento flagelar. A 
peça intermediária apresenta uma parte incompleta formada pelo acúmulo de uma camada amorfa que cerca o axonema 
chamada de bainha fibrosa, o conjunto dessas estruturas é o complexo axonêmico. No espermatozóide de I. iguana a peça 
final flagelar tem o diâmetro reduzido, com o término da bainha fibrosa permanecendo um típico padrão axonêmico de 
microtúbulos (9+2). Uma quantidade grande de citoplasma é observada em torno dos espermatozóides no lúmen dos 
túbulos seminíferos, sugerindo que estes são “espermatozóides imaturos”. As modificações estruturais ocorrem durante a 
sua passagem através do trato reprodutivo. Estas modificações do espermatozóide após ter saído do testículo não foram 
determinadas precisamente ao longo do trato reprodutivo e não são provavelmente essenciais para a mobilidade do 
espermatozóide. Algumas características do trato reprodutivo dos indivíduos machos são mostradas. 
 

PALAVRAS - CHAVE: Reprodução. Lagarto. Ultra-estrutura. Receptáculo seminal. 
 
 
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Biosci. J., Uberlândia, v. 25, n. 2, p. 181-187, Mar./Apr. 2009 

R i d


	 
	INTRODUCTION 
	MATERIAL AND METHODS 
	Specimens of the lizard Iguana iguana were collected in the Pantanal of Mato Grosso do Sul (14o22oS and 53o61oW) Brazil. They were anesthetized with ethyl ether inhalation, the testis and reproductive tract removed, fixed and processed according to routine methods for scanning (SEM) and transmission electron microscopy (TEM) and for light microscopy (LM), stained with hematoxylin-eosin and periodic acid Schiff (PAS). 
	RESULTS AND DISCUSSION 
	ACKNOWLEDGMENTS: We thank the IBAMA for granting permission (proc. # 02014.0077/01-10) to collect a few Pantanal lizard specimens, since Brazilian law protects these animals. This study was supported by FAPESP (proc. # 00/03259-5) and Fapemat (proc. # 0769/2006). 
	REFERENCES