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PHYCOLOGICAL SOCIETY OF AMERICA
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Pugetia fragilissima & Pugetia
firma
Morphology
The morphology of an alga does more than just
differentiate it from its neighbors. It becomes highly specialized,
allowing each algal species to thrive within its unique set
of abiotic and biotic factors. In their book, Seaweed Ecology
and Physiology, Lobban and
Harrison discuss the functional-form model advanced by
Littler and Littler in 1980. The model links the functional
needs of algae, such as photosynthesis, nutrient uptake,
and herbivory survival to the physical form of the weed.
Thus, according to the functional-form model, morphology
is highly specialized to balance each species unique environmental
requirements.
Before the specific function of Pugetia’s
morphology can be understood in the context of its habitat,
it is important to first be familiar with Pugetia’s
basic construction. Although Pugetia firma and Pugetia
fragilissima have a few structural differences, the
morphology and basic cellular functions of these two species
are extraordinarily similar and thus can be generalized together.
For the specific differences in morphology and cellular structure
refer to fragilissima vs.
firma.
Pugetia has an isomorphic life
history, meaning that through all stages of its life cycle
it has the same basis morphology. Its thallus is
erect and almost entirely blade-like with virtually no stipe,
just a small holdfast either centered in the middle of the
blade or at the bottom. The blades are non-calcified, but
instead are cartilaginous and delicate (Murray
and Dixon, 1972). Pugetia firma tends to have
a slightly thicker thallus, whereas Pugetia fragilissima is
a bit finer. Pugetia has
a thin, pigmented cortex and
a thicker multiaxial medulla with
unpigmented isodiametric cells
and small filaments. The cells within cortical tissue appear
to lack organization, however they are arranged into filaments.
Reproductive structures develop in the cortical layer across
the entire blade. To learn about the female reproductive
anatomy in Pugetia fragilissima and Pugetia
firma and its taxonomical significance click here.
Now that a basic description of Pugetia’s
morphology and tissue structure has been described, it is
possible to explore these features in relation to Pugetia’s
unique environmental circumstances in Monterey Bay.
Photosynthesis
Pugetia’s morphology and growth pattern
are essential to ensuring adequate light absorption for
photosynthesis. In 1986, M.E. Hay classified six generalized
thallus morphologies, which absorb light differently (Lobban
and Harrison, 1997).

Pugetia seems to share
some characteristics with two of the thallus morphologies
Hay described. The first is a monolayered flat opaque thallus,
which allows a weed to expose its entire photosynthetic
area to light. When under the canopy of the kelp or on
the shaded vertical faces of rocks, Pugetia takes
on a similar morphology, extending out as if to reach
for light. The second is a multilayered thallus with
translucent blades, which Hays hypothesized, increases
the ratio between thallus area and substrate area and
would provided an advantage in well-lit areas (Lobban
and Harrison, 1997). Translucent blades decrease
self-shading when blades overlap. In this way much more
photosynthetic surface area can be exposed. Pugetia
fragilissima often
utilizes this morphology, having overlapping but translucent
blades at the top of a tubeworm.
Like most red algae, Pugetia has a series of
photosynthetic pigments within its cells, including chlorophyll
a (and d?), carotenoids and phycobiliprotiens, which give
it its beautiful red coloration. The sun’s light
travels at different wavelengths. In general photosynthetic
active radiation is considered to be wavelengths of 350-700nm.
Using a spectrophotometer to measure Pugetia absorption
pattern, it appears that Pugetia’s peak absorption
is at a wavelength of 500nm.
Nutrient Uptake
Unfortunately, nothing is specifically known about Pugetia’s
ability to uptake nutrients. Considering the nutrient rich waters where
it grows, it is unlikely this is a limiting factor, however, Lobban
and Harrison warn against the tendency to generalize the nutritional
requirements of algae. The quantities Pugetia requires may not
be established, but some of the basic nutrients for metabolic processes
are certain to be utilized. These include Cu, Mn, Zn, C, H, O, P, K,
N, S, Ca, Fe, and Mg. Many other nutrients are also needed in some specialized
algae or can be replaced by another element. More work needs to be done
on Pugetia’s nutritional requirements before we know if
it has any such special needs.
In general, a large unbranched thallus and multicellularity
reduce the surface area to volume ratio, thus reducing
photosynthesis and nutrient uptake. The degree of multicellularity
(thickness) of Pugetia is probably not great enough
to make any substantial change in its nutrient uptaking
abilities. However, its pseudoparenchymatous medulla reduces
cytoplasmic contact between the cells of neighboring filaments.
It is believed that the secondary pit connections found
in most red algae solve this problem by creating another
pathway for nutrient exchange (Lobban and Harrison, 1997).
Grazer Susceptibility
Pugetia’s morphology
leaves it relatively susceptible to herbivory. It is not
calcified, nor does it grow as a crust. Pugetia has sacrificed
these lines of defense for fast growth, and large blades
for photosynthesis. To minimize its fitness loss due to
herbivory, Pugetia grows quickly, replacing lost
tissue, and has scattered reproductive structures along
the entire thallus. In this way, Pugetia never
looses its entire reproductive hub to one hungry herbivore.
Additionally, Pugetia may avoid some herbivores
by growing on tubeworms out on the sand flats. For more
on information on this theory go to tubeworms.
Space Constraints
Pugetia grows erect
for its entire life cycle, allowing it to attach to small
areas of substrate, while still expanding its photosynthetic
tissue. With its small holdfast, Pugetia fragilissima can
even grow on the tips of tubeworms. Pugetia’s
cup-like morphology may give it even another competitive
advantage in the battle for open substrate. Some algae with
flat wide growth patterns, such as the form taken by Pugetia on
rocks, may actually block spores from other algae before
they reach the substrate (Lobban
and Harrison, 1997). In this way Pugetia may
guard the substrate around its own holdfast, providing a
possible explanation for why Pugetia firma, when
present, tends to cover the entire rock face, excluding other
species.
Intertidal Stress
Intertidal Pugetia firma and subtidal Pugetia firma have
slightly different morphologies due to the extra stress within the intertidal.
Intertidal Pugetia firma must withstand wave action and exposure to
dry air and direct sunlight, thus it has a slightly thicker thallus with large
medulla cells. The result is a slightly tougher thallus and a lower surface
area to volume ratio reducing the risk of desiccation.
© 2005 Laure Sierra Katz.
You are welcome to use any of my images.
If you have any questions, comments or happy ocean stories
please email me at laurek@stanford.edu
Last updated: Oct. 21, 2005
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