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A . M. G o nzal ez, Chapter 9

CHAPTER 9

Vastago anatomy in selected species of hydrophilic plants

Ana M. Gonzalez

Land plants evolved from photosynthetic aquatic organisms (green algae) and must achieve
a series of characteristics to live on dry land (Raven, 1992). This study analyzed the group of plants
that “decided” to return to their original environment: the aquatic environment. Upon returning to
the water, these plants have to develop adaptations to adapt to the limiting conditions of the
environment, which vary from very light in emerging organs to extreme in submerged organs.

The detailed study of cells and tissues allows for a better understanding of the adaptations
of the plant to a special function and to the end of the various organs in the environment in which it
develops.
This chapter attempts to offer a vision of the microscopic world of hydrophilic plants.
The objective is to analyze and show the anatomy of the stem of some hydrophilic species that
grow in the Iberá macrosystem. The purpose is not to provide an extensive detail of the anatomy of
these plants, but to introduce the reader to the topic and, through the use of microscopic observation
technology, marvel at the beauty that is hidden in our eyes.

Material and Methods

For this work, 28 species of aquatic and marshy plants were selected. Fresh material or fixed
in FAA (70% formaldehyde-alcohol - acetic acid) was used. The numbers of each species correspond
to their respective identification keys.

1.36. Alternanthera philoxeroides 1.30. 1.55. Myriophyllum aquaticum 1.22.

Azolla caroliniana N ymphaea amazonum 1.23.
1.13. Carolinian bomb 1.54. N ymphoides indicates 1.44.
Cephalanthus glabratus 1.5. Oplismenopsis najada 1.35.
Ceratophyllum demersum 1.84. C Pistia stratiotes
yperus giganteus 1.15. 1,193. Polygonum acuminatum
Egeria cobras 1.49. 1.68. Pontederia chordata
Eichhornia azurea 1.40. 1.18. Potamogeton gayi
Eichhornia crassipes 1.103. 1.25. P. ferrugineus
Fuirena Robust 1.42. 1.34. Salvinia biloba
Heteranthera reniformis 1,267. 1.53. Schoenoplectus californicus 3.4.
Hydrocotyle bonariensis 1.41. Utricularia foliosa 3.12. U.
Limnobium spongia 1,134. poconensis
Mayaca sellow iana In the 1.19. Victoria Cruziana
majority of cases, histological sections will be carried out by hand, observation and immediate
recording, given that aquatic plants retain better texture and turgency when fresh, in addition to not
tolerating well the dehydration and assembly process usually carried out in histology. In some
species, definitive preparations could be made, for which the material was dehydrated with Biopur®
dehydrator and clarifyer, embedded in paraffin and subsequently cut with a rotating microtome,
(Gonzalez and Cristóbal, 1997); will be carried out

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transversal cuts (ct) and paradermals of hoja, transversal and longitudinal (cl) of stem. For the
identification of fabrics, double coloring was used: safranin followed by Astra blue (Luque et al.
1996); The determination of crystals was carried out using Lugol's reagent (D'Ambrogio de
Argüeso, 1986) and the identification of crystals through observation with polarized light
(Johansen, 1940).
Observations will be carried out using an optical microscope (OM) that allows the cellular
structure to be seen through histological sections of the material, achieving up to 1000
magnifications (Lám. 9.IC, H). An Olympus BMAX microscope equipped with a clear camera
was used to carry out drawings and diagrams and an Olympus CH30 with which a video camera
connected to a PC equipped with a digitizing card allows the recording of images.
Another tool used for the anatomical study is the barred electron microscope (MEB),
which, unlike the optical microscope, allows observation of up to 300,000 magnifications (Lám.
9.IA, B, DG, IK). For observation under this microscope, the fixed material was dehydrated in an
ascending acetonic series, dried at a critical point in CO2 and metallized with gold-palladium.
Registration is carried out by recording images on floppy disks.
The unit of measurement used for microscopic observations is one micron, equivalent to
one thousandth part of a millimeter (1 µm = 0.001 mm).
For the schemes, the symbol proposed by Metcalfe & Chalk (1950) was used:

Fundamental Clo rénq u ima

Xil ema A erén qui ma
part empalizada

Flo ema Close to lagoon

Col én qui ma E scl erenchyma

Abbreviations used in the figures

alm: almidón, bul: bulliform cells, col: collenchyma, d r: drusen, emp: empalized
chlorenchyma, end : endodermis, esc: sclerenchyma, est: stoma, fib: fibers, f l: phloem, lag:
lagoonous chlorenchyma, lp : protoxylematic lagoon, par: parenchyma, raf: raphidia, sti: styloid;
tan : tannins, x il: xylem; ct: transverse section, cl: longitudinal section.

Anatomy

Submerged organs
Factors conditioning the environment

The plants and plants that live under water must adapt to particular conditions: one is the
reception and capture of sufficient light for photosynthesis, the other is the scarce availability of
carbon dioxide, oxygen and nutrients with lower dissolution in water. However, the problem of
support that must be faced by species that live outside of water has been solved, as support is
provided by the medium (Fahn, 1974; Sculthorpe, 1967).
Hojas
Predominance of the types of leaf forms in the emerged plants: enteras and divided.
The buried hojas are observed in Potamogeton gayi, P. ferrugineus and Egeria snakes, they are
thin to translucent, tapered, linear or lanceolate. This shape is the one that shows the greatest
relationship between surface and volume, which increases the efficiency of absorption of CO2,
O2 and nutrients. The lamina divided into linear to filiform segments characterizes the C abomba
caroliniana, C eratophyllum demersum and Utricularia foliosa. Today's tips are offered smaller

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mechanical resistance to water movement, also benefiting light capture (Sculthorpe, 1967).

The epidermis presents adaptations to the environment as it is: thin cuticle, which facilitates
the passage of substances from the aquatic medium and the presence of chloroplasts,
transforming into photosynthetic tissue. This is directly related to the reduction of the thickness of
the blade, leading to extremes such as the sunken hojas of Egeria cobras
(Lám. 9.3 A, B) where the lamina is reduced to both epidermis, completely lacking mesophyll; In
this case, the epidermis is the only photo-synthesizing tissue in the world. Another characteristic
of the epidermis in submerged leaves is the absence of stomata, only anomocytic stomata were
observed grouped in the abaxial epidermis in the apical zone of the lacinias of C abomba
caroliniana (Lám. 9.2 E; 3 I), Galati ( 1981 ) Consider water-bearing or hydatoid stomata because
they eliminate water. The hojas are generally glabrous, and are often hairless, C eratophyllum
demersum is the only species that poses thorns on the edges of the hojas (Lam. 9.3 E).

The mesophile is homogeneous in all the species analyzed, the absence of differentiation
in clorenchyma in empalizada and lagoonous is due to the variable position of the hoja with
respect to the incident light (Sculthorpe, 1967). The mesophile could be completely absent as in
the lamina of Egeria najas (Lám. 9.3 A, B), reduced to a layer of cells as in the submerged hojas
of C abomba caroliniana (Lám. 9.3 G), Potamogeton gayi (Lám. 9.3 J) y Potamogeton ferrugineus
(Lám. 9.7 M), the presence of several layers of cells and air chambers as in C eratophyllum
demersum (Lám. 9.3 C, D). The submerged leaves generally lack sustaining tissues and the
vascular system is very reduced, only a void of fibers was observed surrounding the main and
lateral haces in the submerged leaves of Potamogeton
gayi and P. ferrugineus, the last species also presents the middle vein with chambers arranged
radially around the vascular haz (Lam. 9.7 L).
Tallos
They are thin and graceful, the main characteristic is the reduction of the number and
lignification of xylematic elements as well as the frequent organization of the vascular system in
a protostele (Arber, 1920). Sustainability is not a problem for submerged plants, which is why
little development of tissues such as collenchyma and sclerenchyma is observed (Fahn, 1974;
Sculthorpe, 1967). This group includes Potamogeton gayi and P.ferrugineus, Egeria najas,
Cabomba caroliniana, Ceratophyllum demersum, Utricularia
foliosa and U. poconensis; Myriophyllum aquaticum is also included although it has an emerging
apical portion.
In short, the stalk presents an epidermis similar to that of sunken leaves (Lam.
9.4). The characteristic of the submerged stems is the formation of extensive air chambers, whose
number and size can vary depending on the size and age of the organ, although their design is
quite specific, in Potamogeton gayi (Lam. 9.3 K) , P. ferrugineus (Lám. 9.7 HJ) and C the
Carolinian bomb (Lám. 9.4 A, B) several cycles of small chambers are present; in Egeria najas
decrease in size inside the cortex (Lam. 9.4 E); in Ceratophyllum demersum, Myriophyllum
aquaticum and in the analyzed species of Utricularia there is a single circle of chambers arranged
radially around the vascular cylinder (Lám. 9.4 C, D, F, G, I, J).

In general, the external portion of the cortex has compact parenchyma (Lám. 9.4 A, B) or
collenchyma with delicate thickenings in its walls (Lám. 9.4 D, col) or sclerenchyma, in small
fascicles as in Potamogeton (Lám. 9.3 K ; 9.7 I, fib). Distributed throughout the cortex are
frequently found cells with abundant starch granules (Lam. 9.4 E, alm), cells

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tanifers (Lám. 9.4 D, tan) and cells with crystals, very abundant in Myriophyllum aquaticum,
especially in the thin partitions that delimit the lagoons (Lám. 9.4 G, cr); The specialized cell or
idioblast that contains these drusen collapses quickly and can only be observed with MEB in
young adults (Lám. 9.4 H).
The vascular system is often represented by a protostele, where the xylem is reduced to
non-lignified, parenchymatous elements, with one or a few central protoxylematic lagoons (Lam.
9.4 D, E, lp), surrounded by phloem elements (Lam. 9.4 D, E, fl). It can also be present
fractionally in two concentric periphloematic haces as in C abomba caroliniana (Lám. 9.4 A) or
in several collateral haces as in Myriophyllum aquaticum (Lám. 9.IV F) or Utricularia foliosa
(Lám. 9.4 J). An endodermis surrounds each hacecillo in C the carolinian pump (Lám. 9.4 A,
end), limits it to the entire vascular cylinder (Lám. 9.4 C, D, E, F, end); This layer regulates the
transfer of mineral ions and water between the cortex and the central cylinder (Fahn, 1974).

Submerged stems and petioles of plants with floating leaf blades In species
such as Victoria cruziana , the petioles must accommodate themselves to maintain large
floating blades, and therefore must extend to prevent them from overlapping one another
(Sculthorpe, 1967). In N ymphoides indicates the stalk has numerous air chambers (Lám. 9.5 J,
K), while the petiole of N ymphaea amazonum and Victoria cruziana is characterized by the
presence of collenchyma and peripheral parenchyma (Lám. 9.IA) and two four large lagoons in
the center surrounded by smaller ones with a specific design for each species (Lam. 9.5 A, B,
D).
The presence of these large air chambers, combined with the fragility of the fabric,
represents a danger of denial, which is contrasted with the existence of diaphragms,
those that are arranged transversally in the air chambers, perpendicular to the direction of the
organ (Arber, 1920; Sculthorpe, 1967). The diaphragms generally present a single layer of cells,
with very small intercellular spaces, if observed in N ymphoides indicates
(Lám. 9.5 O) and in the sunken stalks of Potamogeton ferrugineus (Lám. 9.7 J, K).
Interestingly, diaphragms are absent in the petioles of Nymphaea amazonum and Victoria
cruziana.
The only supporting cells found in these fabrics are sclereid in a star-shaped manner:
astrosclereid, with lignified secondary walls (Lám. 9.5 C, H, I, M, N); in N ymphaea amazonum
and Victoria cruziana they also contain calcium oxalate crystals (Lám.
9.IK), located between the primary wall and the secondary wall (Gaudet, 1960; Ling-Long Kuo-
Huang et al.
2000). Floating organs
Factors conditioning the environment

There is a group of plants that are characterized by having floating leaves, where the
shell is in permanent contact with the water surface. These hojas must maintain stability, adapt
to flotation and resist flooding and immersion caused by water, rain and wind (Sculthorpe, 1967).

Leaf lamina N
ymphoides indica, Nymphaea amazonum and Victoria cruziana form a group with similar
anatomical characteristics. N ymphoides indica and Nymphaea amazonum have suborbicular
or ovate hojas, with a spiked base. The leaves of Victoria cruziana have circular, peltate, strong
blades, with wide and flexible petioles, the edge of the blade is located

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pleated forming a vertical wall 2 to 10 cm high, with an internal margin; It also presents structures
called stomatodes, true orifices through which the bubbles that form beneath the blade can
escape and, if not released, rise apart from the water (Valla & Martin, 1976).

Anatomically, these cells have unique characteristics, as the upper face is exposed to air
and the lower face is in contact with water, resulting in a typically dorsiventral structure (Lám.
9.5 E, F, P). The upper epidermis of the skin is glabrous and has an important cuticle layer,
which helps it repel water; it has abundant stomata (Lam. 9.2 D, G; 9.3 F, I; 9.5 F, P, est). The
lower epidermis may occasionally have stomata, presumably non-functional; A particular
characteristic of this epidermis is the presence of groups of cells that are differentially colored,
called hydropots by Mayr in 1915 (Sculthorpe, 1967), considered structures for the absorption
of ions from the water. In N ymphoides indicates groups of 25-40 cells (Lám. 9.5 L) while in

N ymphaea amazonum is formed by three cells (Lám. 9.5 G). They are also observed in the
submerged remains of both species of Utricularia (Lám. 9.4 K). In Victoria Cruziana, the lower
epidermis presents conspicuous simple uniseriated hairs and multicellular needles, and a large
number of trichomes are also observed on the lower face of the floating leaves of C abomba
caroliniana (Lám. 9.3 F, H).
The mesophyll is made up of a thickened chlorophyll parenchyma, interrupted by numerous
substomatal chambers, with one to four layers of enlarged cells with abundant chloroplasts. On
the lower face of the hoja there is aerenchyma arranged in columns that reach the lower
epidermis, delimiting important lagoons or air chambers (Lám. 9.5 EF, P, emp, lag). In N
ymphoides indica, N ymphaea amazonum and Victoria cruziana
the presence of astrosclereids distributed throughout the mesophyll (Lám. 9.IK;9.5EF, P) stands
out, in N ymphaea additional columnar sclereids are found in the parenchyma and empalized
with branched ends that protrude into the intercellular spaces of the lagoonous parenchyma
(Lam. 9.5 F). The vascular haces are well developed, they have a protoxylem lagoon, the
phloem is well differentiated and is surrounded by parenchymatic veins; The main veins present
a thick cord of collenchyma on the lower epidermis of the hoja (Lám. 9.5 E).

Potamogeton ferrugineus has submerged and floating leaves, these last ones differ in a
greater development of the mesophilus, with a pair of cell layers in the empalized parenchyma
and the lagoonous parenchyma with air chambers, the vascular haces are collaterals and
present sclerenchymatic veins ; Just as in the submerged hut, there are abundant chambers in
the area of the middle vein (Lám. 9.7 N). The lamina has paracytic stomata in the upper
epidermis (Lam. 9.2 L)
The Carolinian bomb has submerged leaves and floating bracts. In the floating bract,
stomata are observed in the adaxial epidermis (Lám. 9.2 F); the mesophyll is reduced to a pair
of layers of cells, the lower one made up of smaller cells (Lám. 9.3 F).
The floating leaves of Limnobium spongia present an adaxial epidermis with epidermal
cells with a wavy contour and numerous paracytic stomata (Lám. 9.2 I). The mesophyll is
dorsiventral, with parenchyma embedded in the adaxial face and aerenchyma in the abaxial
face, which acts as a floater (Lám. 9.6 H, I). Oplismenopsis najada has floating, enveloping
vegetative hobs, which present the mesophyll with air chambers filled with an aerenchyma of
star cells (Lám. 9.9 F, G, H).

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Stems and petioles

Azolla caroliniana, Salvinia biloba, Eichhornia crassipes, E. azurea, Heteranthera
reniformis, Limnobium spongia, Pistia stratiotes are stoloniferous hierbas. The estolones are tall
horizontal ones whose trees unfold new rosettes of hojas. The stems have the characteristics
mentioned above to describe submerged stems, highlighting the presence of air chambers. In
some species, the chambers are periodically obliterated by diaphragms, which vary in the shape
of the cells and the degree of development of intercellular spaces: they are small in the petiole of
Eichhornia crassipes (Lám. 9.8 H, I); in the size of E. azurea the size of the intercellular spaces
increases until the cells have short arms (Lám. 9. 8 D). The stem of Salvinia biloba is very fragile
and presents in the cortex a ring of air chambers with diaphragms whose cells have a particular
arched shape (Lám. 9.7 F; G). Diaphragms will not be found in Pistia stratiotes or Limnobium
spongia.
The presence of idioblasts with _ _ _ _ raphides, styloids, drusen and tannic substances
in the diaphragms and in the aerenchyma that delimits the chambers.

In Heteranthera reniformis the stem presents a central cylinder with numerous hacecillos
included in parenchyma with abundant starch granules, this atactostele is surrounded by an
endodermis and an outer layer formed by cells with thick walls; the cortex presents air chambers
without diaphragms and collateral cells (Lám. 9.10 H); the petiole has an important central
chamber and small chambers on the periphery, all of which have diaphragms; the vascular haces
of the stem and petiole are collaterals and are surrounded by a sclerenchymatic vein (Lam. 9.10
H, I, J).
Oplismenopsis najada is a poácea (grass). It has tall bones with single diaphragms in the
nudos, formed by several layers of star cells (4-6 arms) whose arms delimit intercellular spaces
(Lam. 9.9 K). Emerging organs

Conditioning factors in free-floating plants These plants

must adapt to living in the aquatic and aerial environment simultaneously. It must be able
to float, resist flooding, have a water-repellent surface, etc. (Sculthorpe, 1967).

Hojas arrange these on floating stalks

Azolla caroliniana, Salvinia biloba, Pistia stratiotes, Eichhornia crassipes, E. azurea,
Heteranthera reniformis, Limnobium spongia.
Azolla caroliniana and Salvinia biloba are very reduced plants . B). In Salvinia biloba the
upper face of the leaves presents emergencies with a hollow conoidal base formed by the
epidermis and the chlorophyllous parenchyma, which ends in four arms that remain united at the
apex and trap an air bubble, so that the water cannot penetrate allowing the fleet to be maintained
(Lam. 9.6 C, D, E); Sculthorpe (1967) called them water repellents. Today it has a mesophyll
formed by chlorophyllian parenchyma that delimits large airy lagoons, the lower epidermis has
simple uniseriates and stomata (Lám. 9.6

E, F, est).

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Pistia stratiotes has a very dense coat, formed by simple uniseriate hairs that, according
to Sculthorpe (1967), trap the air in the same way as those of Salvinia biloba (Lam. 9.7 A, E);
These trichomes fall into the old leaves, causing their progressive immersion. The leaf blade of
Pistia stratiotes (Lám. 9.7 A, B) and the leaves of Limnobium spongia (Lám. 9.6 J). It is present
with marked dorsiventrality, with chlorophyllous parenchyma embedded in the upper face,
between this tissue and the lower epidermis there is aerenchyma whose abundant chambers
allow the flotability of the plant. In the mesophyll of Pistia stratiotes, the presence of abundant
calcium oxalate crystals stands out, both in the form of druses and in clusters of roots located in
idioblasts that project into the interior of air chambers (Lam. 9.1 J; 7 B, dr, ra C, D).

The leaves of Eichhornia azurea and E. crassipes have paracytic stomata on both sides
(Lam. 9.2 AB; 9.8 AB); parenchyma empalizada beneath both epidermis, with greater
development in the adaxial face and parenchyma with chambers and diaphragms in the medial zone (Lám.
9.8 A, C, B, F). In the dilated petiole of Eichhornia crassipes the aerenchyma presents large air
chambers forming true flotation organs (Lám. 9.8 G, M), curiously these structures present
scarce support structures, limited to the few lignified elements that provide mechanical support.
It depends mainly on cell turgency, a characteristic that was identified by Siegel (1962) in
Salvinia biloba.
The vascular system of these hojas is formed by abundant vascular cells, the protoxylem
fails to destroy a protoxylematic lagoon and, in general, lacks sustaining elements.

The most interesting characteristic is the presence of inverted haces in the leaves of
Limnobium spongia, Pistia stratiotes and Eichhornia azurea (Lam. 9.8 A), the presence of these
faces would be explained by the compression and lateral expansion of the apical portion of a
ancestral petiole (Arber, 1918,1920; Sculthorpe, 1967).
Conditioning factors in rooted plants Temporary
denial of young plants and leaves that remain under water for a certain period of time
before emerging. Environment of high atmospheric humidity: once the water is dry, the leaves
must control both the loss of water through transpiration and the gas exchange, on which the
supply to the portions of the vegetable that remain submerged depends (Sculthorpe, 1967).

The changes that this group of plants presents are more qualitative than quantitative,
representing a transition to profound modifications that present the submerged plants,
constituting an intermediate state between terrestrial and aquatic plants (Sculthorpe, 1967).

Hojas are arranged on emerging stalks of

Alternanthera philoxeroides, Cephalanthus glabratus, Polygonum acuminatum
(dicotyledons) and C yperus giganteus, Mayaca sellow iana, Fuirena robusta (monocotyledons).

In dicotyledons, the leaves present typical characteristics: lamina with dorsiventral

mesophyll, presence of idioblasts with drusen, empalized parenchyma with three layers of thick
cells, the lagoonous parenchyma generally does not present a very developed system of
intercellular spaces, the leaves they can be hypostomatic or amphistomatic (Lám. 9.10 F, G; 11
B, C, F, G), with scattered, diacytic stomata in Alternanthera philoxeroides (Lám. 9.2 H). The
vascular haces, very differentiated, are surrounded by a

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parenchymatic vein, except in the main veins where they normally present a sclerenchymatic
shell (Lam. 9.11 B, F).
In monocotyledons, leaf structure is characteristic of terrestrial plants.
Mayaca sellow iana is the only species analyzed that has the mesophyll reduced to a layer of
cells and air chambers that run lateral to the medial vein, interrupted by thin diaphragms (Lam.
9.11 I, J). The epidermis presents stomata arranged in longitudinal rows, in Pontederia chordata
the cuticle forms striations (Lám. 9.2 C). The mesophile can be isobilateral with parenchyma
empalizada on both sides as in Pontederia chordata (Lám. 9.9 A) or homogeneous without
differentiation in parenchyma empalizada and lagoonous as in Fuirena robusta, corresponding
to the erect position of these leaves, the epidermis posee stomata on both faces (amphistomatic
hojas), bulliform cells are also observed (Lam. 9.12 K, bull).
Emerging stems and petioles (with submerged portions)
Alternanthera philoxeroides, Cephalanthus glabratus, Polygonum acuminatum, Hydrocotyle
bonariensis (dicotyledons) and C yperus giganteus, Mayaca sellowiana, Fuirena robusta,
Schoenoplectus californicus (monocotyledons).
They can be massive (Lám. 9.12 A, C, H), with aerenchyma (Lám. 9.9 D; 9.10 AC, H; 9.11
A), with hollow medulla (Lám. 9. 10 E; 9.11 D), or hollows with diaphragms (Lám. 9.10 H, J).
In the dicotyledons the cortex presents parenchyma (Lám. 9.1 B; 9.10 E; 9.11 A, D), the
aerenchyma (Lám. 9.10 B, C), the cells can contain grains of starch (Lám. 9.1 CD) or crystals
(Lám 9.1 G). Collenchyma (Lám. 9.10 E; 9.11 A, D, E) and fibers, which may be periphloematic
(Lám. 9.10 E) or be arranged in a continuous ring (Lám. 9.11 A, D), can be seen as supporting
fabrics. The conduction fabrics form a heating element in a single cycle (Lám. 9.10 A, E) or are
available in a continuous cylinder when the stem presents secondary growth (Lám. 9.11 A, D).

In the monocotyledons the stem has abundant aerial chambers, small and without
diaphragms in C yperus giganteus and Fuirena robusta (Lám. 9.12 H, L), with diaphragms in the
stem and petiole of Pontederia chordata (Lám. 9.9 D). The stem of Schoenoplectus californicus
presents chambers occupied with aerenchyma of star cells and periodically distributed diaphragms
(Lám. 9.12 EG); In this species, the leaves are reduced to leaf veins because the outer region of
the stem presents a chlorophyllous parenchyma, which is thickened to carry out photosynthesis
(Lám. 9.12 B). The support is provided by fibers located in the form of subepidermal shells (Lám.
9.12 B, L) and in the veins of the haces (Lám. 9.12 D, I). The vascular system is organized in an
atactostela, with several cycles of hacicillos (Lam. 9.12
B, H). The collateral vascular haces are surrounded by a layer of fibers, presenting the xylem
with a notable protoxylematic lagoon and large metaxylematic vessels (Lam. 9.12 D, I).

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SLIDE 9. 1 Victoria cruziana: A, parenchymatic cells. Hydrocotyle bonariensis: B, chlorophyll

parenchyma; CD, aerenchyma with starch grains; C, MO, D, MEB.
Eichhornia crassipes: E, styloid; F, rafidios. Alternanthera philoxeroides: G, druse.
Heteranthera reniformis: HI, idioblast with raphidia, H, MO; I, MEB. Pistiastratiotes: J, idioblast
with raphidia. N ymphaea amazonum: K, detail of astrosclereid with calcium crystals.

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BLADE 9. 2 E pid ermis. Eichhornia crassipes: AB. Pontederia chordata: C. Victoria cruziana:
D. C abomba caroliniana: E, hoja subergida, F, hoja flotante. Nymphoides indica: G.
Alternanthera philoxeroides: H. L imnobium spongia: I. Azolla caroliniana: J. Hydrocotyle
bonariensis: K. Potamogeton ferrugineus: L. epidermis inferior: A, B, C, E, J, K; ep. top: D, F,
G, H, I, L.

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L AMINE 9. 3 Egeria najas: AB, ct de hoja. C eratophyllum demersum: C, ct de hoja; D, detail

of the sector marked in C; And, sharpen the edge of the blade. Cabomba caroliniana: F, ct of
floating hoja; G, ct hoja subergida; H, lower epidermis floating leaf blade; I, non-functional
stoma in adaxial epidermis. Potamogeton gayi: J, CT hoja; K, ct of the stem.

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BLADE 9. 4 Carolinian c ambob: A, stem ct; B, detail marked on A. Ceratophyllum demersum:

C, stem ct; D, detail of the marked portion in C. Egeria snakes: E, portion of the tall.
Myriophyllum aquaticum: F, stem size G; detail marked in F; H, drusen cells.
Utricularia foliosa: I, ct stem; U.poconensis: J, ct tallo; K, detail of the hydropot in ct.

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BLADE 9. 5 Victoria cruziana: A, petiole ct; B, detail; C, astrosclereid. Nymphaea amazonum: D,

petiole ct; And, today's date; F, detail; G, hydropot cl; HI, astrosclereidas en ct y cl. N ymphoides
indica: J, ct tallo; K, details. L, hydropot in surface view; MN, astrosclereids en ct y cl; O, tall
diaphragm; P, ct today.

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Flora d el Ib erá

L AMINE 9. 6 Azolla caroliniana: A, general view from the cliff; B, emerging leaf lobe.
Salvinia biloba: C, D, E, emergencies from the upper face of the hoja; CD, MEB, D, MO; D, ct
of today; F, trichomes on the lower face of the hoja, G, ct hoja. L imnobium spongia: HI, ct hoja
flotante; J, ct today emergent; H, scheme; I, MEB; J, details.

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SLIDE 9. 7 Pistia stratiotes: A- B, hoja ct; C, air chamber; D, idioblast with raphidia;
And, trichome of the lower epidermis of the today. Salvinia biloba: F, stem ct; G, detail of the
stem labeled in F. Potamogeton ferrugineus: HI, ct stem; J, cl tall in the diaphragm zone; K,
diaphragm cells; LM, ct today submerged; N, ct floating hoja.

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BLADE 9. 8 Eichhornia azurea: AB ct de hoja; , C, hoja diaphragm; D, tall diaphragm; And, style
in today. Eichhornia crassipes: F, ct hoja; G, ct inflated petiole; HI, diaphragm en petiole; JK,
styloids on petiole; L, petiole in the non-inflated portion; M, petiole in the diaphragm zone
showing a tanniferous idioblast.

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BLADE 9. 9 Pontederia chordata: A, ct hoja; B, stomata of the adaxial epidermis; C, today

diaphragm; D, ct petiole; And, petiole diaphragm detail. Oplismenopsis najada: F, ct hoja; G, ct
leaf vain; H, star-shaped aerenchyma of the leaf vein; I, ct tallo; J, haz tallo; K, bare diaphragm.

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BLADE 9. 10 Hydrocotyle bonariensis: A, ct tall; BMEB ; , detail marked in A; C, ct tall with

D, ct today. Alternanthera philoxeroides : E, ct stalk; F, G, ct today. Heteranthera reniformis:
H, ct stem; I, ct petiole; J, detail of the peripheral zone of the petiole with diaphragms in the
chambers.

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A . M. G o nzal ez, Chapter 9

BLADE 9. 11 Cephalanthus glabratus: A, ct tall; B, today's CT; C, detail marked in B.

Polygonum acuminatus: D, ct stem and leaf vein; E, detail of the peripheral zone of the stalk
with tangential collenchyma; F, ct hoja; G, detail marked in F; H, drusen. Mayacasellowiana: I,
ct hoja; J, detail of today and an air chamber with diaphragm.

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BLADE 9. 12 Schoenoplectus californicus: A, ct tall; BC, detail; D, hacecillo; And, detail of the
diaphragm; F, measured at the height of a diaphragm; G, aerenchyma. C yperus giganteus: H,
tall sector; I, hacecillo; J, epidermis. Fuirena robusta: K, ct of hoja at the height of the middle
vein; L, ct peripheral zone of the stem.

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