XI. Magnoliophyta: the Flowering Plants We Come in the End to The

XI. Magnoliophyta: The Flowering Plants

We come in the end to the largest of all vascular plant groups, the flowering plants. There are about 250,000 of them in about 462 families and 40 orders. Three features of flowering plants set them apart from all of the other vascular plants: 1) a folded and sealed leaf called the carpel, which encloses the ovules; 2) a highly reduced female gametophyte, of just eight nuclei in seven cells; and 3) double fertilization, that is the fusion of one sperm with an egg and a second sperm with a neighboring cell of the female gametophyte to yield a nutritive tissue called the endosperm.

A. Apical Meristems

Angiosperms share an unusual kind of meristem with the Gnetophyta (a group we will mention in lecture, but which won't be examined in lab). This meristem, called the tunica-corpus meristem, is well known to biology students because it figures prominently in introductory biology. However, not many divisions of vascular plants have this type of meristem.

1. Examine the prepared slides of Coleus apical meristems available in the lab. The tunica and corpus are visible at the very summit of the stem (the apical dome), along with the primordia (mounds of tissue destined to be leaves and buds).

DIAGRAM the Coleus shoot. Label tunica, corpus, leaf primordia, bud primordia, and leaf trace.

B. Variations in Vegetative Features of the Flowering Plants

1. Many different orders of flowering plants include species that are adapted to xeric (dry) environments. At some point during the lab have a look at the following:

a. Mammillaria: Here is the classic cactus, with leaves reduced to spines and stems fattened to increase water storage capacity.

b. Euphorbia: Here is a plant that has the same adaptations as Mammillaria, but has a completely different flower design. In contrast to the cactus, leaves are still produced by this plant.

c. Lithops: These plants grow in the rough gravel of drylands in Africa: they are called living stones. In this case, the leaves have been highly modified so that they look almost like stones, but still function to store water and gather sunlight energy.

2. Among the many amazing leaf transformations in the flowering plants, those of the passionflower (genus Passiflora) are among the most unbelievable. Look at the passionflower leaves set out on display.

a. First of all, look at the shoots transformed into tendrils that arise from the axils of the leaves. These tendrils attach this plant to the other plants around it in the forest.

b. Now look closely at the petiole of these passionflower leaves. There are a couple of small spherical bodies attached part way out the petiole. These are supposed to be butterfly-egg mimics. A species of butterfly whose larvae eat this species of passionflower is in part diverted from laying eggs on these petioles. It appears to the butterfly that they are already taken by the eggs of another butterfly. Here is a single example of the amazing number of interactions between animals and flowering plants. In fact, one of the most important themes in the study of angiosperms is the interaction with and adaptation to animals.

C. Angiosperm Stem Anatomy

The angiosperm stem has a variable and confusing anatomy, but a there are a few basic features that most have in common.

1. Look at prepared slides of alfalfa (genus Medicago, Fabaceae). Under low power, look for the ring of vascular bundles surrounding a large pith and surrounded by a narrow cortex. Now look closely at the ring under medium power. What you should be able to pick out is a group of vascular bundles disrupted by secondary tissue produced by a vascular cambium. Look for the following critical features:

a. the vessel elements of the xylem, with thick, red-stained secondary walls

b. the sieve tube elements of the phloem, located to the outside of the xylem.

c. to the outside of the sieve tube elements, look for a mass of phloem fibers These have lime-green or reddish staining walls and are irregular in shape and arrangement.

DIAGRAM the stem transverse section. Make a DRAWING of a single vascular bundle.

D. The Structure of Angiosperm Wood (see Fig. 22 for examples of tracheary elements of various woods.)

Wood structure varies quite a bit from family to family among the angiosperms; here we examine the structure of maple (Acer) wood by studying prepared slides. First, identify the three kinds of sections: transverse, radial, and tangential. (Remember, all cells in the transverse section are more or less round. In the radial section the tracheary elements appear long vertically and the parenchyma rays look like brick walls.)

1. In transverse sections, notice that there are large vessel elements distributed randomly through the wood. The remaining cells are either very thick-walled fibers or parenchyma

2. In radial section, study the vessel elements to see how they fit together into vessels. (Remember, the places where the primary wall is missing are called perforations.) The fibers in this section have just a few very narrow pits. Radial rows of parenchyma cells are also visible - these are called vascular rays.

Make a labeled DRAWING of a transverse section of the maple wood.

Fig. 22 Evolutionary trends of tracheary elements and fibers. d, e, primitive long tracheids; f, vessel element with scalariform perforation plate; g,h, vessel elements with simple performation plate; a-c, evolution of fibers. Redrawn from Esau, 1965. Plant Anatomy, Wiley & Sons, Inc., p. 228.

E. The Angiosperm Strobilus

The most familiar feature of angiosperms is the flower. Everybody thinks of flowers as showy structures that attract bees and hummingbirds, but in fact they are extremely diverse in form and function. Many, such as the flowers of grasses and birch trees, are not showy at all. Morphologists agree that, in terms of homology, flowers are strobili, but there are two major schools of thought as to the exact

77 homology of the flower:

1): The flower is a simple strobilus made up of leaves. The lower leaves of the strobilus (the sepals and petals) are without sporangia; the upper leaves bear microsporangia (stamens), and above these, integumented megasporangia (ovules) in folded leaves called carpels.

2): The flower is a compound strobilus. There are leaves below, but the stamens are branch systems, not leaves: microsporangia are borne directly on the ends of stems. The megasporangia are interpreted in the same way as in theory 1.

We will not be able to see enough kinds of flowering-plant flowers to understand and make a choice between these two opinions, but at least you can have the two ideas in mind as you look

1. Begin by reviewing basic flower structure with a dissection of Tradescantia. Pick a flower or group of flowers from the plant in lab and use a probe to dissect one under the dissecting microscope. You should see:

a. Most prominent: a whorl of bright purple petals that function in the attraction of pollinators.

b. Lowermost: a whorl of small, green sepals that functioned in protecting the buds (look for the sepals on the buds of Tradescantia flowers on the plant). These sepals also function to support the open petals. Petals and sepals together are called the perianth.

c. Just inside the petals: a whorl of six stamens. In the flowering plants stamens are made up of two parts, a stalk (called the filament) and a synangium (called the anther), usually of four connate sporangia. The filaments of Tradescantia have long, delicate hairs - you can see cytoplasmic streaming in these hairs if you mount one of your filaments on a slide and look at it under the microscope. Be sure to check to see that your Tradescantia stamens have four sporangia per synangium.

d. In the center of the flower is the pistil. Three distinct regions of the pistil can be recognized: the broadest part (at the base) is the ovary, which is where the ovules are located. Above the ovary is the slender style, and at the

78 tip of the style is the stigma. In Tradescantia the stigma has a beaded appearance under the dissecting 'scope.

DIAGRAM a flower of Tradescantia

2. Perianth Transformations

Now look at a series of flowers to get an idea of the kinds of transformations possible among angiosperm flowers. The basic themes here are fusion and reduction. Look for connation, adnation, and loss of floral structures.

a. Go out to look at the flowers of Magnolia on the green with the rest of your group. These flowers are quite different from Tradescantia:

- The perianth parts are all alike, and there are many of them.

-The stamens are numerous and they are flat and somewhat leaflike (hence the idea that stamens are homologous with leaves)

- The pistils are numerous, and they are attached along an axis.

All agree that Magnolia is a primitive angiosperm - the argument is as to how primitive. You should be able to gather that this flower looks a lot more like a strobilus than does Tradescantia, mostly because the axis of the flower is elongate like other strobili we have studied.

Another thing that's important to think about: here the pistils are many, and each has a single stigma. The conclusion from many observations like this one is that these pistils are simple - no connation is involved in their history, whereas the pistil of Tradescantia is taken to be compound, the product of connation of three simple pistils like those of Magnolia.

b. Now, back in the lab, dissect a flower of Browallia under the dissecting scope. Look for two kinds of fusion, connation and adnation. In this flower the petals are connate into a trumpet- like structure: we infer that there were five petals involved in the fusion because there are five lobes to the trumpet. Now, open up the trumpet with a probe. Inside, you should find that the stamens are adnate to the narrow tube of the trumpet.

79 F. Germination and Growth of Pollen (Start at beginning of lab)

In this part of the lab, you conduct a simple experiment to demonstrate the growth of the pollen grain (immature male gametophyte) once it has reached the stigma, since pollen grains will germinate on agar supplemented only with sugar. The rest of the nutrients needed for growth are stored inside of the pollen grain.

1. Your teaching fellow will put a drop of sugar agar on a clean slide for you.

2. Wait a minute for the agar to cool and solidify. (Hot agar can cook the pollen -- and you get no results from cooked pollen.)

3. Flowers with pollen will be in covered dishes in the lab. Lightly tap the stamens of these flowers over your sugar agar. Check with a microscope to be sure pollen is in your agar. Mark your slide with your initials and the time using a wax pencil or marker. Finally, put the slide in one of the large covered glass dishes in the lab to keep the agar culture moist. Use a probe to dislodge pollen if it doesn't fall out when you tap.

4. After 15 minutes place one drop of 16% sucrose solution on top of your agar with pollen. Return the slide to the covered dish and check it once in a while for pollen tubes. Once you can see good pollen-tube development under the microscope (say a pollen tube ten times as long as wide) add a small drop of toluidine blue stain and then a coverslip to the culture. In another ten minutes, study the pollen tube closely to try to locate the nuclei in the tube.

In real life, the pollen tube grows down a canal in the style and into the ovary, with the egg inside the ovule as its goal. Many pollen grains can be growing down the canal at once, and there is a competition among pollen grains that appears to be controlled by the pistil. In other words, the female parent can choose among male candidates in the style!

5. In slide #10 of the Lilium development slides, you can see the style of a pistil covered with pollen. Some of the pollen has germinated and begun to grow, just as the pollen in your sugar agar did.

After one hour, count the number of pollen grains, and the number of germinated pollen grains on each slide. (Count approximately 30 grains on each slide). Evaluate your data to determine if there were any differences in

80 germination between the two treatments.

*** The following chemicals have been shown to affect pollen tube germination. All also contain 16% sucrose.

Auxin – can increase pollen tube germination at low concentrations (1x10-6 M) but can be inhibitory at high concentrations (1x10-4M). Involved in cell (acid) growth pathway.

Boric Acid – has been shown to increase germination. Involved in cell wall synthesis.

Cobalt (as CoCl) – inhibits germination. Blocks calcium channels.

Gibberellic Acid – can increase germination at most concentrations.

G. Fruits and Seeds

The pistil is in fact easiest to study when the ovules have been fertilized and embryos have developed. The ripened pistil is called a fruit, and as usual the ripened ovule is called a seed. We have already looked at the simple pistil of Magnolia and the compound pistil of Tradescantia. Now we need to think a bit more about the homology of the simple pistil itself.

Pea and bean pods provide a vivid example of simple pistils that look like folded leaves with seeds inside of them. In fact, the most popular idea for the homology of the carpel is that it is a folded and sealed leaf (sealing is a new kind of fusion, fusion of a plant structure to itself, like sealing an envelope). Strong arguments based on anatomy and morphology of angiosperms assumed to be among the most primitive alive support the homology of the simple pistil with a single folded and sealed leaf. This hypothetical ovule-bearing leaf has come to be known as a carpel.

1. Consider the fruits of pea or bean available in the lab.

a. Dissect open a pea or bean pod. Notice that this ripened pistil has a single withered style at the tip and a withered perianth at the base. You may even find a withered stamen or two. Inside, the seeds are arranged in a single row along one side of the cavity.

b. Make a transverse thin section of a pea or bean pod and stain it for a while in phloroglucinol.

Make a DIAGRAM of the pod both 1) opened up to reveal the interior and 2) in transverse section.

2. Now consider the cherry tomato fruits we have set out. These provide an excellent example of fruit structure derived from a compound pistil (that is, a pistil comprising two or more fused carpels). Make a transverse section of a cherry tomato. How many carpels are present in this compound pistil?

Make a DIAGRAM of the transverse section of the tomato fruit near your pea or bean pod diagram, so that you can compare the two.

3. Variations and Dispersal Syndromes

Finally, consider the small collection of fruits in the lab. Fruits are in fact adapted to specific kinds of dispersal. Fruit and seed structure are often dramatically altered to fit specific dispersal needs. In spite of these dramatic alterations, you can usually discern the basic homology of the simple or compound pistils from which the fruit was derived. For each of the following fruits, identify: 1) What is ovary wall? 2) What is a seed? 3) How many carpels make up the fruit? 4) Is the pistil simple or compound?

a. peanut b. olive c. orange d. banana e. strawberry f. kiwi