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The Shoot
Angiosperm
structure, and that of vascular plants in general, is the result of
meristematic activity. Meristems are centers of mitotic cell
division that produce tissue of the plant body in a manner similar to
laying bricks. The largest and oldest living things
are vascular plants and this is due to the nature of the meristem.
Apical or terminal meristems produce root and shoot tissue and
these - in essence - are the two 'organ systems' of the vascular plant.
Structures and associated botanical terminology discussed here for both
vegetative and reproductive expressions relate, mostly, to the product of
the apical or terminal shoot meristem and this pattern of activity defines
a fundamental 'framework' of structural organization (see OSU
Botany 300, source of the image [left], for more detail).
As indicated by both size and longevity of some vascular plants, the
apical meristem is indeterminate. Its activity of producing
cells, or laying the bricks of the plant body, can continue indefinitely.
The apical shoot meristem also has the capability to produce secondary
centers of meristematic activity, lateral meristems. The
lateral meristems provide a structural groundplan for mapping the
angiosperm shoot. The position of lateral or axillary meristems, as
either lateral buds or secondary shoots and often subtended (positioned
beneath) by a leaf, is known as a node. Shoot tissue that
extends between these points is knowned as the internode.
Aside
from the lateral meristem, another secondary meristematic product of the
shoot meristem - the vascular cambium - is significant. This
is a column of dividing cells that produces vascular tissue (xylem and
phloem). Since meristematic activity of the vascular cambium
varies according to available resources and conditions during the growing
season, its product - wood - often takes the form of 'annual rings'.
Angiosperms with an active vascular cambium are 'woody' and those
taking a tree-like form (single stem greater than 4-5 m in height) are arborescent
whereas woody angiosperms with a shrub-like aspect (more than 1 shoot,
less than 4-5 m) are fruticose.
Most angiosperms lack a
vascular cambium, i.e., they are non-woody herbs or herbaceous.
Both woody and herbaceous angiosperms can show indeterminate apical
meristem activity and a perennial life style. Herbaceous
perennials living in temperate or seasonable parts of the world sacrifice
'above ground' tissue during the difficult season but often retain dormant
meristems and photosynthate in a tough, underground structure known as a caudex.
However, most herbaceous flowering plants are annuals in that they
show an annual cycle of seed germination, vegetative growth, and
reproduction with the next generation passing the 'difficult' annual
period (Winter, dry season) within the seed or fruit. Summer
annuals germinate in the Spring and pass the Winter as seed whereas Winter
annuals germinate in the Fall, flower in the Spring, and pass the dry
Summer season as either seed or fruit. Many key players in the
remarkable Texas Spring flora are Winter annuals that are in full
vegetative growth during December and January. Some angiosperms
dedicate a full growing season to vegetative growth, sequester
photosynthate - often in underground structures - through the difficult
season and invest this in a reproductive effort at the start of the second
year. These biannuals are often found at the market (carrot,
onion, etc.) because we harvest the saved resource prior to its 'intended'
use by the plant. Many herbaceous angiosperm genera of central Texas
include species with varying 'life styles' (annual, binnual, perennial)
and this can be a factor or 'key character' for identification.
Thus, it is important to insure that the sample collected includes a
'below ground' component. Also, terms are discrete and biological
variation can be continuous and, as a result, difficult to define using
specific terms. Consequently, 'bridging' terms can be applied.
The term suffrutescent, for instance, denotes an herbaceous perennial
that shows some woody tendencies, usually at the base.
The stem 'caul-'
refers to the primary product of the terminal meristem, the shoot.
Caulescent denotes a 'normal' shoot with nodes and discernable
internodes. Acaulescent, on the other hand, descibes a
situation where the internodes are reduced and, as a result, this shoot is
compressed to the point that the plant looks 'stemless', such as head
lettuce (Lactuca - Asteraceae) and cabbage (Brassica -
Brassicaceae). Terms relating to various adaptive permutations of
the shoot include:
Erect - 'normal'
growth toward the sun
Prostrate - a horizontal shoot, flush to the ground
Decumbent - a horizontal shoot, but not flush to the ground - the
tips point upward
Caespitose - producing clustered, multiple shoots forming tufts or
cushions
Scandent - twining, climbing - vine-like
Scapose - acaulescent but producing an erect, leafless flowering
stalk (scape) - Taraxacum officinale
Function modifications of
the shoot include:
Rhizome -
horizontal, elongate and underground, often thick and fleshy (herbaceous
perennial are sometimes rhizomatous see: Cyperus
- Cyperaceae) Tuber
- thick and fleshy and usually underground, like a rhizome, but compressed
- via reduction of the internodes - and enlarged as a water/photosynthate
storage structure (Potato - Solanum
- Solanaceae)
Corm - vertical,
compressed and underground, often thick and fleshy (herbaceous perennial) Bulb
- acaulescent with expanded, fleshy leaf bases (Onion - Allium
- Liliaceae) Stolon
- horizontal, above-ground, often rooting at the nodes - strawberry (Fragaria
- Rosaceae) and St. Augustine grass (Stenotaphrum
- Poaceae) are stoloniferous.
Thorn - a hard,
sharp-pointed modified shoot (Honey locust - Gleditsia
- Caesalpiniaceae)
Branches of deciduous, woody plants in the Winter show both action of the
apical meristem and quite a few useful key characters. Keys are
often available for woody plants in 'Winter condition' for a given flora.
Each Fall the apical meristem is enclosed and protected by a set of
terminal bud scales. These scales fall away each Spring, leaving a
ring of bud scale scars on the branch. Thus, the annual product of a
given terminal meristem, nodes and internodes, can be tracked by examining
the portion of the branch that lies between the terminal bud and the first
set of bud scales scars encountered. Other features, such as the
leaf scar and pattern of vascular bundle scars within the leaf scar, are
used for identification in these keys.
Another feature of the
shoot that can be used as a key character is the nature of the central
tissue or pith.
The pith is normally
present as soft, uniform tissue (continuous), but this can be
intersected by discs of more dense tissue (diaphragmed) with, in
some cases, the 'pithy' soft tissue absent (chambered).
The Leaf
Supported by the stem, leaves are the primary
site for photosynthesis in most flowering plants. Their structure
and position on the stem also provides a rich suite of taxonomic
characters. The fossil record suggests that the first angiosperms
had simple, alternate, entire leaves. These
terms relate to: structure, position on the stem (or phyllotaxy),
and leaf margin.
STRUCTURE:
Simple - blade a
single, undivided structure (see compound) - example
Compound - blade divided into substructures (leaflet), each
with its own stalk (petiolule)
Pinnately Compound -
leaflets arranged along a main axis (rachis) - example
'Even' = no terminal leaflet, 'Odd' with a terminal leaflet
Bipinnately Compound - as above but leaflets compounded - example
Palmately Compound - leaflets with petiolules attached to a
single point - example
Phyllotaxy
Alternate - a single
leaf at the node
Opposite - two leaves at the node, each usually subtending
(immediately below and close to) a lateral bud or shoot
Whorled or verticillate - more than two leaves at the node
Rank - when applied to leaves, denotes rows of leaves along
the shoot
Decussate - four ranked or opposite leaves alternating at right
angles to those above and below
Cauline - leaves are those associated with the central shoot
Fascicled - closely clustered or grouped |