NECTAR AND NECTAR SECRETION.

The great Swedish botanist, Linnaeus, nearly two hundred years ago,

basing his classification of plants on their flowers, found it

necessary to name and account for all of the parts of a flower. In many

cases he found structures that were neither sepals, petals, stamens nor

pistils, and as these contained, or were wet with, a sweet fluid, he

gave this the fanciful name “nectar”--the drink of the gods, and called

the parts of the flower that produced or contained it, “nectaries.”

As these nectaries were different from stamens and pistils, which

Linnaeus recognized as the sexual organs of flowers, though they

are sometimes connected with them, and as they were different from

ordinary sepals and petals, though sometimes connected with them,

they presented something of a question to the men of that day, who

were curious to know what the parts of a plant really are and what

they do. For this reason the study of nectaries became something of

a popular diversion for a generation or two; and a general idea that

they are organs for secreting sugar became established; not necessarily

an idea of secretion, though, for just as animals excrete various

organic substances that are by-products, or waste, from some of their

functions, so it was thought by some students that the sugar or nectar

might really be an excreted waste or surplus rather than a substance

secreted because it is to become useful to the plant.

[Illustration: Fig. 98. Sprengel’s title page.

Das

entdeckte Geheimniſs

der

NATUR

im Bau und in der Befruchtung

der

Blumen

von

Christian Konrad Sprengel,

Mit 25 Kupfertafeln.

Berlin, 1793.

bei Friedrich Vieweg dem æltern.]

Toward the end of the eighteenth century a German rector, Sprengel,

who seems to have found in nature a good deal of inspiration that he

failed to “put over,” noticed that the petals of the common German

wild geranium were fringed with hairs at their bases. That was in the

day before men believed in evolution, but when they did believe in a

purposeful creation. Sprengel was convinced that an all-wise Creator

would not have made a single hair in vain, and he set about discovering

what these hairs were for, much as a sensible person, seeing the

governor on an engine today would try to find out what it is for. Below

the break between the petals, he found a nectar gland, producing its

sugary fluid; and he saw the hairs would prevent the nectar from being

diluted or washed away by rain or dew. This brought him back to the

original question--what nectaries and nectar are for. He got his answer

to this by watching the plant and seeing that bees visited the flowers

and removed the nectar as what might be called the raw materials of the

honey industry.

In Sprengel’s day the general impression was not only that things have

been created just as we find them, but created for our own ultimate

good. So Sprengel found an answer in discovering that the hair fringe

of the geranium petals protects the nectar of the flowers and so

preserves it for bees to use in manufacturing honey for our breakfast

table.

When you stop to think about it, Sprengel could hardly have had the

curiosity to study out his geranium question to an answer without

being spurred to look at other flowers to see if they might not have

something interesting of the same sort to offer. He yielded to the

impulse to look at other flowers, and he found his geranium to be a

very drab specimen compared with some of the irregular and painted

flowers that he studied out in the same way. He must have felt no

common pride when, in 1793, he published the results of his studies,

with simple but effective illustrations, under a title that meant

revealing to mankind the newly discovered secret of nature in the

structure and fertilization of flowers.

But Sprengel seems not to have been the sort of man to whom such an

answer really was an answer, and he looked further. It does not seem

to have taken him long to see that while gathering their own store

of honey, and obviously without consciousness that they were doing

anything else, the bees became dusted with pollen from geranium stamens

and rubbed it off on geranium stigmas while going their rounds of the

flowers. This conclusion evidently answered two questions--what the

hairs are for, and what nectar is for.

It is not necessary to walk down Michigan Boulevard on a windy day to

realize that we belong to an imitative race. The corner grocery and the

drug store show it as well as the windows of milliners and dressmakers,

shoe shops and news stands, or as the sights that issue from a barber

shop in a college town.

Fashions run in fads and interests quite as much as in dress. Linnaeus

was a great botanist; perhaps none has been greater. He not only

reduced a chaotic science to order, but interested men in its study to

a remarkable extent. It is rather unfairly charged against him that

because his service was somewhat one-sided, those whose interest he

awakened were extremely one-sided, in that they did not see or care for

much in botany beyond finding, describing and classifying new plants.

This was well enough worth doing; it is not finished yet, and will not

be finished for many years to come; but it had become so fascinating

and workable through the genius of the Swedish master that his

followers seized on it with eagerness, and it was a long time before a

mind of original habits and impulses broke loose from the train.

The man who possessed this originality was Darwin, the promulgator of

the now universally accepted idea of organic evolution. To him has been

ascribed the introduction of a new teleology into natural science,

recognizing that structures and functions are because they are or have

been of use--not of use to man necessarily, though man may turn them to

account--but to their possessor.

[Illustration: Fig. 99. Ant acacia.]

This was Sprengel’s conclusion as to the nectar of geranium flowers,

which he found led to their fertilization. The essential difference

between his way of seeing it and Darwin’s is that he thought the entire

mechanism has been specially made by the Creator as a means to an end,

while Darwin saw in it the gradual modification of earlier structures

because the new were helpful in the struggle of life and their

possessors for this reason were likely to survive and pass them on to

their offspring.

There is a German country saying that the honeybee was forbidden the

red clover because she didn’t keep Sunday. Beekeepers know that her

tongue is a little too short for the honey tube of the red clover

flower and that she doesn’t waste time in trying to get what is out

of her reach. They know, too, that some races of honeybees really can

suck the red clover nectar because they have longer tongues, and if

beekeepers ever want to do it they can probably set an expert plant

breeder to work at breeding a race of red clover with a tube short

enough so that even the German honeybee can get at its nectar. Natural

evolution hasn’t done this. Where red clover is at home bumblebees are

found, and bumblebees have no difficulty in reaching its nectar much as

hawk-moths get that of a moon flower, which is far beyond the reach of

any kind of bee. But in the South Seas, where there are no long-tongued

bees, red clover finds itself as unable to set seed as the German

honeybee is to get at its nectar. Bee and flower have evolved together,

where both are at home, into a harmony of structure that is helpful to

them both.

Nothing was more suggestive to Darwin in his search for evidences of

evolution, or modification through descent, than this sort of harmony

of structure and habit in flowers and insects; and one of his earliest

and most effective books in bringing his views to the comprehending

notice of others was one dealing with the mutual relations between

those freaks in flowers, the orchids, and their insect visitors.

For Sprengel’s teleology, Sprengel’s explanation of nectar as a means

of securing fertilization was sufficient. For Darwin’s teleology, it

carried another question: =Why?= The geranium flower has both stamens

and pistil, standing in its middle. The one might fertilize the other

just as well as not, apparently; and yet this does not happen, for

the pollen-bearing anthers of the stamens drop off before the stigmas

of the pistils come to maturity. The same thing may be seen on any

single-flowered geranium in a bay window or a greenhouse or a summer

window box or flower bed, only this geranium does not belong to the

genus =Geranium= of the botanists, but to the related African genus

=Pelargonium=.

Looking for a further reason, Darwin saw a step further into the

mystery when he found that these and many other flowers that ought to

get on without any help are as dependent upon insects through their

own failure to bring pollen and stigma together as those are in which

stamens and pistils are borne in separate flowers--often on separate

plants. To him, nectar and its attendants--flower fragrance, color,

variegation, bizarre shape, long tubes, nectar guards of hairs or some

other structure--meant what they had meant to Sprengel, fertilization

through insect aid; but they meant something more, fertilization of one

flower by pollen from another flower--crossing.

And still the questions multiply. Why do not all flowers have stamens

and pistil side by side? Why, when they have this structure, do

they not time the maturity of these essential parts so as to secure

effective functioning without all the nectar machinery? In other words,

=why= is crossing so commonly necessitated and provided for?

Science of every kind has been advanced by three methods--reasoning,

observation, experimentation. Sprengel’s answer was reached by the

first two; the new answer sought by Darwin was to be obtained through

the third. For eleven years he put the question direct to the plants

themselves, fertilizing them by their own pollen, crossing them,

raising and requestioning their offspring. More and stronger progeny

from crossing was the answer.

The popularity that Linnaeus had given to characterizing and

classifying living things, was transferred by Darwin to studying their

structure and doings. Sprengel’s idea fell upon barren soil; Darwin’s

was cultivated with care and skill.

Two men--Mueller, a German, and Delpino, an Italian, stand out most

prominently among a multitude who observed and wrote and pictured

the marvels of flower and insect harmonies for a generation. All did

excellent work in furnishing details and corroborations; but Darwin

had answered the question as to the what and the why of the nectar of

flowers.

But there is nectar that is not produced in flowers. Look at the

queer spots in the angles between the veins on the under side of a

catalpa leaf, when it is young; or at the little goblets on the stalk

of a cherry or peach or snowball leaf, or at the pin-head spots on a

trumpet-creeper or paeony calyx, and you may see glands there that

secrete a sweet fluid. Bees may not care for it, but wasps or ants do.

The cotton plant has such sweet nectar glands on the outside of the

cluster of bracts about each blossom, and on the back of its leaves.

In a very few cases such “extrafloral” nectar serves the same purpose

as that within the flowers; but generally it does not lead to

fertilization. Delpino called the nectar that leads to fertilization

“nuptial” nectar, and the other “extranuptial.”

In the seventies of the last century an English mining engineer,

Belt, well known in the ore regions of Colorado, was marooned by his

profession on a mining property in Nicaragua. Using his eyes took the

place with him of tennis, or of dissipation, which is the white man’s

bane in the tropics. He saw that a certain kind of ants cut the leaves

of trees into bits which they take into their nests, and that roses and

other introduced plants fared hard with these leaf cutters, unless they

were protected by aromatic oils, as various kinds of citrus leaves are,

or in some other way.

Belt did not fail to notice that ants visited extrafloral nectaries in

numbers. In the case of those on some acacias he found the ants very

pugnacious. I confess that in Guatemala I have preferred, myself, to

go around a bush or a grove of such acacias with their ant guards. As

with Sprengel’s geranium hairs, these nectaries unfolded question after

question.

In Belt’s case, the tips of the acacia leaflets ripen also into little

fruit-like bodies that the ants gather and take into their nests;

and they make these nests in the stipules that flank each leaf and

sometimes are shaped like a pair of small buffalo horns. It is an

interesting undertaking to get the ant census of an acacia twig of this

sort. The danger may not be as great, but it is as real and perhaps

as painful as in taking the census of a mountain valley noted for

moonshine traffic.

Belt drew the conclusion that extranuptial nectar, sometimes

supplemented by solid food and shelter, is of use to the plant that

provides it by maintaining a bodyguard of ants on plants that otherwise

would be defoliated and injured by leaf cutters or grazing animals,

much as Sprengel and Darwin found an explanation of nuptial nectar in

the benefit of insect pollinization of the flowers.

This is the simple story of nectar, simply told, as it has been

seen by observing and thinking men. But it is not a story free from

complications. Our blue violets rarely set fruit from their showy

nectar-bearing flowers, but their main reliance for seed is on

flowers produced below the leaves, and these do not open, but are

self-fertilized. The beautiful =Poinsettia=, with its brilliant red

bracts and large cups overflowing with thick nectar, does not fruit

in West Indian gardens any more than it does in our greenhouses at

Christmas time. And irresistibly pugnacious as the acacia ants are,

those that visit our paeonies and cassias and other plants do not

usually more than protest mildly if we molest the plants that they are

on.

Are the explanations of Sprengel and Darwin, and of Belt wrong? No

other that are at all satisfactory have been offered.

When one stops to think of it, the secretion of nectar is an unusual

phenomenon. Sugar is made within plants and it does not leak from them

unless they have been injured. The sugar beet takes various substances

out of the soil water, but it does not permit the passage of sugar

into the soil water. And yet nectar, essentially sugar, is passed out

of the plant, within which it was manufactured. This is because it is

secreted--or excreted--through specialized glands. Everyone who grows

plants in a bay window has seen young clover or grass leaves with a

drop of water on their tips at some time or other. A few grains of

bird seed in a flower pot covered by a pane of glass will show this as

quickly as the seedlings come up.

These drops pass out finally through pores; but they are drops of water

and not of nectar. If we can imagine a gland behind such a water pore,

secreting sugar--letting it really get out of the cells with or into

the water, we can picture a nectar gland. Such glands occur in some

flowers. Some botanists believe that extranuptial nectar glands were

originally water glands that have acquired the habit of secreting sugar.

This habit is a very unusual and a very peculiar one. It is not really

understood except as it may be connected with usefulness to the

plant. If this usefulness is not indirect, in the ways suggested by

Darwin and Belt or otherwise, it must be direct. Water glands relieve

over-pressure when absorption is high and evaporation low; in some of

the calla family the water even spurts from the tips of the leaves at

times. But sugar is not like water, taken in in quantity and to spare;

it is manufactured, and in the case of nectar glands it is manufactured

where it is secreted. Nobody has yet suggested any physiological

function of plants calling for sugar safety valves situated in

the queer positions occupied by extranuptial nectar glands, and no

satisfactory direct physiological explanation of the nuptial glands has

been suggested.

The actual status of nectar in botanical science is about this: When it

is produced in flowers, and in some cases when it is outside of them,

but near them, it demonstrably serves to secure cross-pollinization

through the aid of insects, or humming birds and their like, when

the flowers are long, tubular and red, as in the trumpet creeper,

the trumpet honeysuckle and the scarlet salvia. It is then “nuptial

nectar.” When it does not serve the plant in this way and so is

“extranuptial,” it occurs in the neighborhood of the flowers, as in

cotton, sweet potato, trumpet creeper and paeony, where it attracts

numbers of ants, which are often very pugnacious, and to the extent of

their activities it prevents injury to the flower buds and flowers,

especially in their early stages; or it occurs on developing leaves

during the period of their greatest need. More rarely, as in the

acacias, the leaves continue to secrete it through the season, so that

those that are mature add to the protection of the younger leaves and

the flowers and the young fruit.

That neither of these functions is served in exceptional cases, and

that some flowers rely on the wind for effective pollination, or have

lapsed into self-fertilization; or that really pugnacious ants do not

commonly frequent the extranuptial glands of most plants in temperate

regions, and that some plants get along very well without such help,

mark questions that will continue to stimulate observation and

experiment. But nothing now known of the physiology of plants offers

an alternative explanation for that which connects nectar with either

pollination or defense; and until such an explanation can be found

nectar will continue to be regarded as connected indirectly with these

services through insect or bird relations.--William Trelease, Botanist,

University of Illinois. American Bee Journal, Dec., 1919. (See also

Physiology of Nectar Secretion).

NEGUNDO, see Box Elder.

NEVADA--Honey Sources of.

Much of Nevada is mountain and desert country. Alfalfa is the principal

source of surplus honey, though sweet clover also yields. There are

numerous minor sources, as willows along the streams and many of the

desert plants.

NEW BRUNSWICK--Honey Sources of.

In New Brunswick the principal sources of nectar are alsike and white

clover, fireweed, goldenrod and aster. Willows and maples stimulate

early brood rearing.--F. W. L. Sladen.

NEW HAMPSHIRE--Honey Sources of.

The honey flora of New Hampshire is very similar to that of other

New England States. In spring, willows, maples, fruit blossoms and

dandelions keep the bees busy in preparation for the main flow, which

comes from the clovers. Goldenrod is also important. Sumac, laurel,

blueberries and asters are worthy of mention, also.

NEW JERSEY--Honey Sources of.

Plants from which nectar is gathered in less than surplus quantities:

Maples, mid-March-early April.

Peach early April.

Pear, mid-April.

Apple, late April early May.

Willows, late April.

Dandelion, early May.

Wild strawberry, May.

Lupine, May.

Raspberry, May.

Grape, late May-early June.

Huckleberry, blueberry, late May-late June.

Persimmon, mid-June-late June.

Vervain, late June-early September.

Virginia creeper, late June-late July.

Milkweed, silkweed, July.

False indigo, July.

Button bush, July.

Burdock, July-October.

Tree of Heaven, pride of China, July.

Catnip, July.

Motherwort, August.

Smartweed, heartsease, late August-mid-September.

Horsemint, August-September.

Boneset, mid-August-September.

Surplus honey plants:

Crimson clover, mid-May.

Locust, May 20-June 1.

Tulip-poplar, May 26-June 10.

Poison ivy, mid-May-mid-June.

Holly, late May-late June.

Mountain laurel, late May-late June.

Sheep laurel, late May-late June.

Swedish clover, alsike clover, June 1-July 10.

White clover, early June-mid-July.

Dogbane, Indian hemp, early June-late August.

Basswood, linden, late June-early July.

California privet, mid-July-late July.

Sumac, mid-June-mid-July.

White sweet clover, June-November.

Cranberry, June 15-August 15.

August flower, soap bush, sweet pepper bush, late July-late August.

Rose mallow, swamp mallow, late July-early September.

Burdock, July-November.

Spanish needle, mid-August into October.

Heartsease, smartweed or blackheart, August-September.

Heath aster, white aster or St. Michaelmas daisy, late

August-mid-October.

Bushy goldenrod, late August-mid-October.

Buckwheat, early August.

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