Jm. Dannenhoffer et Rf. Evert, DEVELOPMENT OF THE VASCULAR SYSTEM IN THE LEAF OF BARLEY (HORDEUM-VULGARE L), International journal of plant sciences, 155(2), 1994, pp. 143-157
The development of the vascular system in the leaf of barley, Hordeum
vulgare L., was studied from its inception to maturity. Young primordi
a were examined with the scanning electron microscope, and distinctive
early morphological stages of primordium development were correlated
with the initiation of the longitudinal procambial strands. All longit
udinal procambial strands originate in isolation, without continuity w
ith the previously existing stem vasculature. The first-formed strands
, which have large-bundle anatomy in both blade and sheath of the matu
re leaf, arise in the disk of insertion (the stem segment associated w
ith the primordium) and from there develop acropetally into the free p
ortion of the primordium and basipetally into the stem below. Subseque
ntly formed strands arise in the primordium. These strands are interpo
lated between older strands across the width of the primordium in a pr
ecise order, and they appear to develop both acropetally and basipetal
ly. By the time the young leaf is 4 mm long, all of its longitudinal p
rocambial strands have been initiated; in addition, the ligule has bee
n initiated, delimiting blade from sheath. The first protophloem and p
rotoxylem elements of the large strands arise in the disk of insertion
near the site of initiation of each strand, the protophloem in advanc
e of the protoxylem. From the site of initiation, the protophloem and
protoxylem develop acropetally into the primordium and basipetally int
o the stem, where, at a later stage of development, they interconnect
with phloem and xylem in the nodal region of the leaf two nodes below.
During further leaf growth, three successive regions can be identifie
d within the blade: (1) a distal, mature region in which elongation ha
s ceased and metaphloem and metaxylem elements are differentiating bas
ipetally in all strands, (2) a region in which elongation is occurring
and protophloem and protoxylem elements are mature or nearly mature b
ut in which metaphloem and metaxylem elements have not yet begun to di
fferentiate, (3) a region in which cell division is occurring and some
protophloem and protoxylem elements are still differentiating.