Hemichordata
Ulrich Welsch, Ludwig-Maximilians University of Munich, Munich, Germany
The hemichordates are marine invertebrate organisms forming a phylum within the
‘Deuterostomia’; they comprise two classes, Pterobranchia and Enteropneusta.
Basic Design
Common to all hemichordates is the division of their
bodies into three regions: prosoma, mesosoma and
metasoma. The prosoma is a locomotory preoral lobe,
the short mesosoma contains important parts of the
nervous system and is often called the ‘collar’, and the
metasoma is the trunk. This tripartite division is not only
external but also internal, with three sets of coelomic
chambers.
All hemichordates have a specific heart/kidney organ in
their preoral lobe, which is supported by an epithelial
outgrowth of the buccal cavity called the stomochord. The
heart pumps blood into a cluster of convoluted capillaries
(glomerulus) covered by podocytes which are typical for
sites of ultrafiltration. The ultrafiltrate collects in the
coelomic cavity of the prosoma, the protocoel, and is
passed to the exterior through a short nephridial (coelo-
mic) duct. A basically similar heart/kidney organ is to be
found in echinoderms and also in the chordates. The
stomochord is a notochord-like structure, responsible for
the name Hemichordata (‘half-chordates’). At the begin-
ning of the trunk a pharynx with openings to the exterior,
occurs; these are the gill slits or ‘pharyngeal clefts’, another
character shared with true chordates. Essential parts of the
nervous system are located dorsally, again as in chordates;
in the Pterobranchia the collar contains a dorsal cerebral
ganglion (Figure 1a), and in the Enteropneusta the collar
has a neuronal dorsal cord (Figure 1b).
Pterobranchia
The Pterobranchia are the more primitive hemichordate
group. At present about 20 species are known, most of
which belong to the genus Cephalodiscus, the other genera
being Rhabdopleura and the little known Atubaria.
The Pterobranchia are small (1–5 mm long), colonial,
tube-dwelling epibenthic animals. The tubes in which the
animals live form a branching network, called the
coenecium, consisting of prostrate and vertical tubes which
can grow in Cephalodiscus into 30 cm large masses. The
coenecium is secreted from glandular cells of the preoral
lobe (oral shield). The individual animals of the colony live
in the vertical tubes and are termed zooids. The trunk of the
zooids always forms a long muscular stalk. In Rhabdo-
ENCYCLOPEDIA OF LIFE SCIENCES © 2001, John Wiley & Sons, Ltd. www.els.net
Introductory article
Article Contents
. Basic Design
. Pterobranchia
. Enteropneusta
pleura the stalks of all zooids in a colony join a common
thread-like stolon which runs through the prostrate tubes.
In Cephalodiscus the stalk forms a terminal adhesive disc
which produces several buds that grow into adult animals.
All animals that originate from one adhesive disc form a
small colony. The animals move around in and outside
their tube on their flexible oral shield. The collar bears 2–18
slender arms with pinnately arranged tentacles, function-
ing as a lophophore. The sac-like trunk has in its initial part
a pair of lateral gill pores, which are only absent in
Rhabdopleura. The pterobranchs are filter feeders, using
their ciliated arms and tentacles to create a water current, in
which food particles are transported into the ventrally
located mouth; excess water is eliminated from the pharynx
through its gill pores. Digestion occurs in the simple
stomach.
Pterobranchs reproduce asexually (by budding) or
sexually. In Rhabdopleura later developmental stages are
brooded in the tubes by females. The short-lived (1 or 2
days) and uniformly ciliated larvae are lecithotropic and
settle on a suitable substratum where they gradually
transform into a zooid.
Pterobranchia occur in deep seas and in shallow waters
of tropical, temperate and cold seas. They are particularly
abundant in deep Antarctic waters. The fossil record of
Rhabdopleura-like species goes back to the Middle
Cambrian and the highly diverse exclusively fossil group
Graptolithina (Cambrian to Carboniferous) is presumed
to be closely related to rhabdopleurids.
Enteropneusta
The Enteropneusta (‘acorn worms’) are benthic animals
that are most common in shallow waters, but single species
have been found in deep seas and at hydrothermal vents.
There are about 70 species, many of which live in mucus-
lined burrows in sand or mud or under stones in coarse
gravel or in seaweed meadows.
Most species are 10–40 cm long, but some species reach 1
to 2.5 m in length. The prosoma forms a slender or oval
proboscis which is mobile and contractile. The collar does
not bear arms and tentacles, though their function is taken
over in several species by the proboscis; its ciliated surface
can transport food particles, trapped in a mucus film, into
the mouth at the base of the collar. However, burrowing
1
 Hemichordata

Anus

Lophophores
Ganglion
Gill slit
Gonad Mesosoma
Pericardium
Stomochord
Prosoma
Intestinal Coelomic
tract pore
Epidermis
Metasoma
Mouth Muscle
Stalk Glomerulus Glandular
Bud cells

(a) Collar cord

Coelomic
pore
Gonads
Hepatic Pericardium
pouches Glomerulus
Intestinal Muscle
tract

Prosoma
Metasoma Epidermis
Stomochord
Anus Mouth
Mesosoma
(b) Gill slits

Figure 1 A diagrammatic drawing of the body plan of the hemichordata, (a) Pterobranchia, (b) Enteropneusta. Due to their considerable length, the stalk
of the Pterobranchia and the metasoma of Enteropneusta have not been drawn in complete length. Numerous epidermal cells are of glandular nature as
indicated by fine stippling.

species are predominantly deposit feeders, taking up mud
or sand and digesting out the organic components. At the
ventral base of the proboscis a preoral ciliary organ occurs,
which may play a role in directing the water stream into the
mouth. The epidermis of the base of the proboscis is richly
endowed with nervous tissue (fan-shaped organ). The
collar contains dorsally the tube-shaped collar cord, a
component of the nervous system resembling (both in
development and structure) the chordate neural cord. The
stomochord, which extends forward from the mouth cavity
into the proboscis has a microscopic structure which
resembles that of the vertebrate notochord; it is built up by
large vacuolated epithelial cells and is surrounded by a
fibrous sheath. The enteropneust trunk is the longest part
of the body and is divided into three sections: the
branchiogenital, the hepatic and the intestinal region.
The branchiogenital region (the pharynx) is characterized
by two dorsal rows of a few to about 200 U-shaped gill slits.
Presumably the gill slits serve to remove excess water taken
up through the mouth and also to exchange respiratory
gases. (Laterally in this region the gonads form two genital
ridges.) The hepatic area is marked by pouches of the
intestine with digestive and resorptive epithelium. The
intestine runs straight to the anus at the end of the trunk. In
burrowing species typical faecal casts can be found at low
tide on the surface of exposed sands.
The sexes are separate in the acorn worms, eggs and
sperm being released into the water where fertilization
takes place. Most species have long-lived planktotrophic
larvae, the tornaria larvae, which have complex systems of
ciliary bands at their surface. These larvae bear a striking
resemblance to some of the echinoderm larvae, probably
indicating a close phylogenetic relationship between
Hemichordata and Echinodermata.
Further Reading
Balser EJ and Ruppert EE (1990) Structure, ultrastructure, and function
of the preoral heart-kidney in Saccoglossus kowalewskii (Hemichor-

2

data, Enteropneusta) including new data on the stomochord. Acta
Zoologica 71: 235–249.
Benito J and Pardos F (1997) Hemichordata. In: Harrison FW (ed.)
Microscopic Anatomy of Invertebrates, vol. 15, pp. 15–101. New York:
Wiley Liss.
Dilly PN, Welsch U and Rehkämper G (1986) On the fine structure of the
alimentary tract of Cephalodiscus gracilis (Pterobranchia, Hemichor-
data). Acta Zoologica 67: 87–95.
Hemichordata
Ruppert EE and Barnes RD (1994) Hemichordata. In: Invertebrate
Zoology, 6th edn, pp. 865–877. Philadelphia, PA: Saunders.
Nielsen C (1995) Pterobranchia (chap. 47); Enteropneusta (chap. 50). In:
Animal Evolution. Oxford: Oxford University Press.
Welsch U (1984) Hemichordata. In: Biology of the Integument, vol. 1, pp.
790–799. Heidelberg, Germany: Springer Verlag.
3
Hemichordates:
Development
John Gerhart, University of California Berkeley, California, USA
Christopher Lowe, University of Chicago, Chicago, Illinois, USA
Marc Kirschner, Harvard Medical School, Boston, Massachusetts, USA
Hemichordates resemble chordates in important aspects of their development anatomy
and physiology. Within the hemichordate phylum are two main subgroups, the
enteropneusts, of a few hundred species, and the pterobranchs, of 20 or so species.
Comparison of hemichordates and chordates allows deductions about the common
ancestor from which they both evolved.
Introduction
The phylum of bilateral animals closest to our chordate
phylum is that of the hemichordates. Study of their
development and organization has long been expected to
illuminate the evolution of chordates. Within the hemi-
chordate phylum are two main subgroups, the enter-
opneusts, of a few hundred species, and the pterobranchs,
of 20 or so species. All are marine animals having gill slits
and a three-part body (prosome, mesosome and meta-
some). Each part contains a coelom or coelom pair see
Hyman, 1959, and Benito and Pardos, 1997, for anatom-
ical descriptions). Enteropneusts, with worm-like bodies as
exemplified in Figure 1, live in burrows or under objects,
some at great depths, but most in shallow water. The pro-
some is a conical, muscular proboscis (acorn-shaped in
some species, hence the name ‘acorn worm’) used for bur-
rowing and for sweeping suspended food particles towards
the mouth by the motion of cilia. The mesosome serves as a
nerve-rich, muscular collar that covers and uncovers the
mouth during food sorting, and the elongated metasome
bears gill slits anteriorly and the long gut posteriorly. A
wide mouth perforates the prosome–mesosome boundary
on one side, designated ventral. Some enteropneusts (the
Harrimaniids) develop directly from the egg to the juvenile,
and others (the Ptychoderids and Spengeliids) develop
indirectly from the egg to a tornaria larva, and then a
juvenile. Hemichordate-like fossils have been dated to the
mid-Cambrian.
Pterobranchs bear little resemblance to enteropneusts.
They are small (1 mm), sessile, tube-dwelling, direct-
developing, deep-sea animals, living in colonies of thou-
sands of individuals, connected by thin bridges. The
three-part body has one pair of gill slits, a post-anal stalk
and ciliated feeding tentacles extending from the mesosome
near the mouth. Pterobranch-like graptolites, now extinct,
were abundant in the Ordovician and Silurian periods.
ENCYCLOPEDIA OF LIFE SCIENCES & 2007, John Wiley & Sons, Ltd. www.els.net
Advanced article
. Introduction
Article Contents
. Development of Saccoglossus kowalevskii, an
Enteropneust Hemichordate
. The Fate Map of S. kowalevskii
. Chordate-like Traits of Hemichordates
. The Domain Map of Hemichordate Embryos
. The Tornaria Larva
doi: 10.1002/9780470015902.a0004110
Gill slits suggest the affinity of hemichordates to chor-
dates, and the tornaria larva suggests an affinity to
echinoderms. Recent phylogenies based on compared de-
oxyribonucleic acid (DNA) sequences (Figure 2) place
hemichordates as the sister-group of echinoderms, which
develop pentameric organization as adults, very different
from hemichordates. The two phyla together (the ‘ambu-
lacraria’) constitute the sister group of chordates (Brom-
ham and Degnan, 1999; Cameron et al., 2000). These three
phyla constitute the supertaxon of deuterostomes, so
called because the mouth (-stome) develops from a sec-
ondary (deutero-) site in the embryo, not from the primary
(proto-) site of the blastopore used by embryos of many
other phyla (the protostomes). Deuterostomes share a
common ancestor from which the three phyla evolved.
Development of Saccoglossus
kowalevskii, an Enteropneust
Hemichordate
William Bateson, (1884, 1885, 1886) who later invented the
word ‘genetics’, first reported the developmental stages as
shown in Figure 3. This species inhabits sand and mud of the
intertidal zone of the Atlantic coast of North America from
Maine to Florida. Adults are 4–10 in long. Gravid females
bear grey-coloured ovaries along the sides of the meta-
some, and males bear orange coloured testes. Arthur and
Laura Colwin described the stages more fully from 1952 to
1960, making a simple fate map, and assessing the capacity
of separated embryonic cells to develop (Colwin and Col-
win, 1950, 1951). We began developmental studies in 2000.
Hemichordates in general are not well studied, and this
species is better known than any other.
1
 Hemichordates: Development
Although the natural triggers of spawning are unknown,
ripe females can be induced to release eggs in the laboratory
by brief warming (278C, 6 h; Colwin and Colwin, 1962;
Lowe et al., 2004). The unfertilized egg is uniformly grey in
colour, 0.3 mm in diameter and moderately yolky. It is
arrested at first meiotic metaphase, with the spindle close to
the surface at a point, the animal pole, where polar bodies
will form after fertilization. Eggs can be fertilized in the
laboratory by sperm released into sea water when one dis-
sects a small fragment of testis. The fertilized egg develops
to a hatching juvenile in 6 days at 228C. Major events
(Figure 3) are as follows:
Day 1: After fertilization, the egg changes shape at the
times of first and second polar body extrusion (20 and
45 min), becoming pear-shaped in one direction and then
Proboscis
Mouth
Tail
Anus
Figure 1 Hemichordate adult (left panel) and juvenile (right panel). The
acorn worm, Saccoglossus kowalevskii, is shown. On the left, bar 1 cm; and
on the right bar, 1 mm.
Protostomes
Lophotrochozoa
Ecdysozoa
Figure 2 Phylogeny of the hemichordates, based on DNA sequence comparisons. Note that hemichordates, echinoderms and chordates share a
common ancestor, the deuterostome ancestor.
2
the opposite. Cytoplasmic materials segregate to the an-
imal pole and the opposite vegetal pole at these times.
Cleavage first occurs at 130 min, and then every 30 min,
until at 12 h, a thin-walled (single layer) spherical blastula
is formed of a few thousand cells. The blastula flattens at
the vegetal pole, and by 18 h the vegetal hemisphere inverts
within the animal hemisphere, to produce a two-layered
cup-shaped early gastrula embryo. The rim of the cup,
which is the blastopore, then contracts in the next 8 h.
Day 2: By 24–30 h, the blastopore closes to a point, the
future posterior end, and the embryo is spherical and dou-
ble layered. A ring of cilia, the telotroch, forms near the
blastopore, and the embryo rotates rapidly within the spa-
cious fertilization envelope. The embryo then elongates,
and two circumferential grooves form, the earlier marking
the prosome–mesosome boundary, and the later, the
Collar mesosome–metasome boundary.
Gill Day 3: A short lengthwise groove appears in the meso-
slit
some (collar) on the dorsal side and neural ectoderm sub-
merges in a neurulation-like movement. Internally,
Pharynx pouches of mesoderm protrude anteriorly and bilaterally
from the inner layer of cells. The pouches soon detach and
Gut expand as coelomic cavities, and the endoderm of the inner
layer seals together to form the tubular gut.
Day 4: The first gill slit pair forms, and the mouth per-
forates the surface. An apical tuft of cilia extends anteri-
orly. The metasome elongates on the dorsal side between
the telotroch and the anus (blastopore), bending this part
ventrally. The telotroch remains close to the anus on the
Deuterostomes
Chordates Hemichordates
(Saccoglossus kowalevskii)
Cephalochordates
Echinoderms
Ptychoderidae
Harrimaniidae
Urochordates
Pterobranchs
Xenoturbella
Spengeliidae
Vertebrates
Enteropneusts
Ancestor of deuterostomes
Ancestor of bilateral animals
 Hemichordates: Development

40 min 130 min 3h 5h 7h 18 h

Unfertilized Fertilized Blastula Early

egg egg gastrula

24 h 28 h 30 h 38 h 3 days 4 days 6 days

Late Telotroch Anterior Posterior Gill slit Mouth

gastrula formed groove groove forms perforates Hatching

Figure 3 Developmental stages of S. kowalevskii. Drawings are modified from the 1884 publication of William Bateson, and they have been arranged in
order and labelled. The temperature of development is approximately 228C.

ventral side. The body contracts intermittently, due to

newly differentiated muscles, and the embryo continues to Animal pole
rotate in the fertilization envelope. and polar bodies
Day 5: The mouth perforates. The embryo elongates Future
further, and bends further. The ventral postanal tail l
bilatera
appears as a small stub. plane
Future anterior epidermis
Day 6: The juvenile hatches from the fertilization enve- and nervous system
lope and swims a few hours by its cilia, attracted by light.
When it contacts sand grains as a substratum, it attaches
avidly and burrows between them, making a micro-burrow. Future posterior epidermis
The newly hatched juvenile (1 mm long) possesses most and nervous system
aspects of adult organization, though sexually immature.
It begins feeding after a week. It has one pair of gill slits at Future gut, muscles,
hatching and then adds more in succession during growth; coeloms
Vegetal pole
the adult has 40–80 pairs. The sticky, contractile, postanal
extension, or ‘tail’, of the juvenile is used to attach to ob-
jects and it retracts when startled. This tail is shed before
adulthood in direct developing enteropneusts; it is appar- Figure 4 Fate map of S. kowalevskii. A schematic drawing of the 8-celled
ently never formed in indirect developing enteropneusts. embryo, which the Colwins (1953) marked with a Nile blue spot at various
places. Embryos were then allowed to develop to the juvenile to see what
(Pterobranchs have a posterior stalk, possibly related to
tissues and organs were blue. The initial and final positions of the marks are
the tail.) related by the fate map.

quartet, those originally near the animal quartet, remain

The Fate Map of S. kowalevskii
To make a fate map, Colwin and Colwin (1951) marked 2, 4
and 8 cell embryos with spots of Nile blue dye, and then
recorded the spots’ locations in the subsequent juveniles.
Thus, they could determine the developmental fate of each
part of the cleaving egg. Progeny of the animal quartet of
cells of the 8-cell stage remain outside during gastrulation
and become anterior ectoderm of the juvenile, including
the nervous system (Figure 4). Some progeny of the vegetal
outside and become the posterior ectoderm of the juvenile,
while progeny originally near the vegetal pole move inside
during gastrulation and become mesoderm and endoderm.
Our recent studies show that progeny of the vegetal-most
tier of cells of the 32-cell embryo are those that move inside.
They also found that the first cleavage plane coincides
with the bilateral plane of the juvenile. Second cleavage,
then, separates pairs of cells that will develop to dorsal or
ventral parts of the juvenile. However, they could not
predict whether a particular pair was to become dorsal or

3
 Hemichordates: Development

ventral. Events determining the dorsal–ventral axis remain
unknown.
Chordate-like Traits of Hemichordates
Four traits identify chordates: the notochord, dorsal hol-
low nerve cord, gill slits and postanal tail. Because Bateson
thought hemichordates possessed most of these, he placed
them in the chordate phylum. Other researchers were du-
bious, and by the mid-twentieth century, hemichordates
were relegated to their own phylum, the Hemichordata
(‘half-chordates’) (Hyman, 1952). The following sections
update the comparisons of hemichordates and chordates,
and the deductions about their common ancestor.
Gill slits: In hemichordates, these develop like the gill
slits of amphioxus (a nonvertebrate chordate), have similar
gill bars and express similar genes. The deuterostome an-
cestor probably had gill slits, which were just carried for-
ward in both lineages (though lost in echinoderms).
Postanal tail: The ventral extension of the enteropneust
juvenile may be homologous, as a body region, to the dor-
sal chordate tail (Burdon-Jones, 1955) since both express
similar genes (hox10–13). The deuterostome ancestor
probably had such a body region, though it may not have
functioned for swimming.
Notochord: The stomochord of hemichordates, which is
a short rod of vacuolated cells protruding into the prosome
from the anterior end of the gut, has been likened to the
chordate lineage. Further study of the hemichordate nerv-
ous system may help to resolve the ambiguity.
In summary, at least two of the four traits of chordates
may have predated them, since the traits are shared by
hemichordates.
The Domain Map of Hemichordate
Embryos
Although hemichordates and chordates differ greatly in
their overt anatomy, they are similar in a fundamental and
covert ‘molecular’ anatomy, namely, the spatial domains
in which certain genes are expressed in the embryo. These
genes encode transcription factors and signalling proteins
of importance for early development. As summarized in
Figure 5, many genes expressed in the developing forebrain
of chordate embryos are expressed in the developing pro-
some of hemichordates. Other genes, expressed in the de-
veloping midbrain of chordates, are expressed in the
mesosome and anterior metasome of hemichordates, back
to the level of the first gill slit. Yet others, expressed in the
chordate hindbrain and spinal cord, are expressed in the
hemichordate metasome posterior to the first gill slit.
Finally, as noted above, genes expressed in the chordate
tail are expressed in the hemichordate postanal tail (Lowe
et al., 2003). Thus, the anterior–posterior organization of

lim1/5
dmbx

chordate notochord. Evidence weighs against homology.

barH
dbx
otp

irx
tll

It is anterior to the gill slits in hemichordates whereas the

hox9/10
notochord is posterior to the gill slits in chordates. Genes wnt1
pax6

pax2

hox7/8
nkx2.1

emx
otx
en

hox1
hox3
hox4
expressed distinctively in the chordate notochord are not
bf-1
six3

gbx
vax

cdx
dlx
tbr

ptf
rx

expressed in the stomochord, which instead expresses

genes like those of the chordate prechordal endomeso- Meso- 1st gill slit
some Anus
derm. At present, a notochord-like precursor cannot be Prosome Metasome
attributed to the deuterostome ancestor. The notochord Hox11/13a
may have originated within the chordate lineage. Mouth
Hox11/13b
The nervous system: Hemichordates have a diffuse intra- Hemichordate Hox11/13c
epidermal nerve net, not a centralized system (Bullock, Mid-
1946, 1965; Knight-Jones, 1952). Nerves arise throughout brain Hindbrain Spinal cord
the ectoderm, interspersed with epidermal cells. Some ax- Hindbrain spinal cord
Fore-
ons connect locally in a basal mat, and others extend for brain
long distances, gathering in bundles at the dorsal and ven-
tral midlines and serving as through-conduction axon
tracts (Cameron and Mackie, 1996). The tracts were ini- Chordate
tially called nerve cords, suggestive of homology to the
Figure 5 The similar domain map of hemichordates (above) and
chordate dorsal hollow nerve cord. However, they are not chordates (below). Each domain is the specific region of the embryo where
neurogenic since they lack neuronal cell bodies, and they a particular gene is expressed. These genes encode proteins of importance
probably process little or no information since they lack in the animal’s development. Lines are drawn vertically to set off anterior,
interneurons. Whether the deuterostome ancestor had a middle and posterior sections of the map, to show that the genes expressed
diffuse or centralized nervous system is debatable. Argu- in the chordate forebrain are expressed in the hemichordate prosome, that
genes expressed in the chordate midbrain are expressed in the
ments can be made on both sides (Holland, 2003). If cen- hemichordate mesosome and anterior metasome back to the first gill slit,
tralized, hemichordates and echinoderms would have lost and that genes expressed in the chordate hindbrain and spinal chord are
it. If diffuse, centralization would have occurred in the expressed in the hemichordate metasome posterior to the first gill slit.

4

members of these two phyla is very similar in terms of the
map of these domains. The deuterostome ancestor pre-
sumably had such a map.
In the dorsal–ventral dimension, the two groups are less
similar. Recalling the differences mentioned previously,
hemichordates lack a central nervous system and a noto-
chord, two dorsal landmarks of chordates. Nonetheless,
both groups appear to organize other aspects of dorsal–
ventral development according to a polarized distribution
of signalling proteins, Bmp and Chordin, which are pro-
duced in the early embryo on opposite midlines (Lowe
et al., 2006). Between these midlines, a graded Bmp distri-
bution is set up. In both groups the gill slits and heart
develop on the Bmp side, whereas body wall muscle de-
velops on the Chordin side. Numerous genes are also ex-
pressed in dorsal–ventral domains in relation to Bmp and
Chordin. These similarities can be attributed to the de-
uterostome ancestor. Several authors have suggested that
the Bmp-Chordin polarity dates back even to the ancestor
of all bilateral animals.
However, the mouth differs in its location. It is on the
Chordin side in hemichordates and the Bmp side in chor-
dates. Since the mouth defines the ventral side of the body,
by scientific convention, hemichordates and chordates
have an inverse dorsal–ventral orientation. By an old and
plausible explanation, ancestral chordates inverted the
whole body and evolved a new mouth opposite the old (De
Robertis and Sasai, 1996). Evidence is still incomplete for
or against this ‘inversion hypothesis’. According to the
findings cited above, hemichordates would be uninverted,
as would be the animals of all other bilateral phyla except
chordates (Nübler-Jung and Arendt, 1996).
The Tornaria Larva
Ptychoderid and Spengeliid enteropneusts develop indi-
rectly to the adult via a tornaria larva that resembles the
larvae of some echinoderms. Since chordates lack such a
larva, the deuterostome ancestor may or may not have
possessed one. The early steps of indirect development (e.g.
Ptychodera flava) resemble those of S. kowalevskii, though
starting from a smaller egg (0.12 mm). After the gastrula
stage, the embryo develops a protocoel cavity and pore, but
the development of gill slits, body musculature and aspects
of the nervous system are delayed until metamorphosis.
Like direct developers, the embryo develops an apical tuft
and a telotroch of cilia, but unlike them, it adds elaborate
ciliated bands around the mouth, at which time it is des-
ignated a ‘tornaria’ (Latin, torno, to turn). These bands
serve the larva’s intake of food particles in the plankton
(Figure 6). After several weeks or months of feeding and
growth, the larva initiates metamorphosis; feeding cilia
disappear, the prosome–mesosome boundary constricts,
the posterior parts lengthen and gill slits start to develop
Hemichordates: Development
Apical tuft
Protocoel
Protocoel
pore
Mouth
Ciliary
band Stomach
Telotroch Intestine
Figure 6 The tornaria larva. A schematic diagram of the larva of
Balanoglossus claverigus, side view. The thick black lines indicate rows of
cilia, one around the mouth for feeding and one near the anus for
movement. Depending on the species, these rows may be more elaborate
with protruding loops.
(Hyman, 1952; Nakajima et al., 2004). The elongating ju-
venile burrows into the intertidal sand and there completes
metamorphosis, including differentiation of the posterior
coeloms and gut.
The map of gene expression domains found in the direct-
developing S. kowalevskii embryo and juvenile is incom-
plete in the P. flava tornaria. Many domains do not appear
until metamorphosis, in keeping with the delayed develop-
ment of many juvenile parts. The fate map of P. flava re-
sembles that of S. kowalevskii in the gross allocation of
parts (Henry et al., 2001), when fates are scored in the
hatched larva. The long larval period has prohibited re-
searchers from identifying the embryonic precursors of
adult structures.
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5
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Van der Horst CJ (1927–1939) Hemichordata. In: Bronn HG (ed.)
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