Journal of Heredity 5: 255-267 (1914)
Pueblo Indian Maize Breeding
Varieties Specially Adapted to Arid Regions Developed by Hopis and Navajos
Their Work Not Sufficiently Appreciated—Probably Much Yet to be Learned from Them.

G. N. Collins
Botanist, Bureau of Plant Industry, U. S. Department of Agriculture,
Washington, D. C.

AMONG the many gifts that the white man received from the American Indian, maize is undoubtedly the most valuable. In the mythology of the Indian this cereal occupies a central position as the most important gift of the gods.

The power of minute observation commonly possessed by primitive people is highly developed in the agricultural Indian tribes of the Southwest. These Indians spend much of their time in their maize fields and it is only natural that the plant which supplies their principal food should come to be known in great detail. Every character of the plant, every operation in its cultivation and every stage in its growth are observed by the Indian with a minute attention in which he is seldom equalled by his white brother.

The ceremonies and care with which maize is cultivated by the Indians of the Southwest are well illustrated in a series of articles on "Zuni Breadstuff," by the late Frank Cushing, who showed that the importance of selection, care of seed, and cross-breeding, though not appreciated as such, are all given careful and conscious consideration by the Zunis.

According to Cushing, "In each cornroom or granary of Zuni, are preserved carefully, four objects; an ear of yellow corn full to the tip of perfect kernels, called a yä'-po-to; an ear of white corn which has resulted from the intergrowth of two or more ears within a single husk-fold, called, from its disproportionate breadth and flatness, a mi'-k'iap-pan-ne; a moderately large normal ear of corn which has been dipped by a Seed-Priest in the waters of the great sacred Salt Lake far south of Zuni (Las Salinas of New Mexico), and a bunch of unbroken corn-soot. The latter two objects are laid side by side on the floor in the middle of the corn-room, and upon them also side by side, usually connected by a bandage of cotton filaments, the yä'-po-to and the mi'-k'iap-pan-ne.

"The significance of all this is both interesting and poetic. The corn-soot is held to symbolize the 'generation of life,' the salted and sanctified ear of corn, the material given by the gods and prepared by man, as the means whereby generated life is sustained, and finally, both these are regarded as the resting place or couch of the Father and Mother of corn-crops or seed; the yä'-po-to being the male, the mi'-k'iap-pan-ne, the female.

THE PARTS OF A SEEDLING MAIZE PLANT
This diagram will furnish the non-botanical reader with the names needed to understand the story of Pueblo Indian maize breeding. (Fig. 5.)

EARS SELECTED WHILE GROWING.

"In a field of growing maize the owner selects such hills as give promise of speediest maturity. These receive his special care. No sooner have a few ears ripened on them than he picks the most perfect, as well as a bunch of soot from some neighboring stalk, and tenderly carries them home in his arms. Arrived at the entrance-way of his house he calls to the women within:

" 'We come!'
" 'Ah! How come ye?' say they.
" 'Together, happily,' he replies.
" 'Then enter ye!' calls out the chorus of women's voices, whereupon the man goes slowly in. One of the women beckons his attention to the 'sitting place,' which, in this instance, 'is a decorated basket-tray in the center of the room. Thither he proceeds and places, one by one, the ears of corn in the tray—using care that they shall all point eastward—and lays the bunch of soot over them. The women of the house flock to the mantel whereon stands the family bowl of prayer-meal, each taking a pinch of the sacred substance, while one of their number, the 'corn-matron,' hastens away to the granary, and carefully lifting the ya'-po-to and mi-k'iap-pan-ne, brings them forth. As she nears the tray, she says, across the objects in her hands (addressing the new corn), 'My children, how be ye these many days?' Then the new corn is supposed to reply through the voices of the other women, now gathered near, 'Happily, our old ones, happily!' With this the corn matron deposits her burden on the new bunch of soot, and all present say little prayers significant of the occasion and setting forth their wishes for 'age of life, happy fortune and the health of strength born of the food of maize.' This ceremonial is called the 'Meeting of the Children.' and is performed in commemoration of the return of the lost corn maidens under the guidance of Pai-a-tu-ma, and their welcome by the Seed-Priests of ancient Zuni.

"With the closing of the prayers, the right hand of each worshipper is passed gently over the tray—while scattering prayer-meal—and breathed from. The corn-matron then returns to the granary, bearing both the old corn and the new.. She replaces the old bunch of soot with the new, laying the former away with the fresh cars of corn, and returning the yä'-po-to and mi'-k'iap-pan-ne to their resting place.

EARS CAREFULLY SORTED.


Seedling of Hopi maize planted eight inches underground. The first permanent roots can be seen sticking out seven and one-half inches above the seed. (Fig. 6.)

"When all the harvest has been gathered, dried, sorted and corded up, around and over the 'Father and Mother' in the corn-room, the ceremonial interrupted at the beginning is resumed. While the corn is being classified as to color and grade, the finest ears of each kind are selected and laid aside. These, and the ears of 'new corn' are together laid along the outer edge of the corn-pile. Next morning the corn-matron takes a basket tray—perhaps the same one used before, or at least one like it—and goes to the door of the corn-room. Here she slips off her left moccasin, then enters. As she passes the threshold she looks around as though she were about to address a group of waiting friends and exclaims:

" 'My mothers and children, how be ye and how have you come unto the morning?' —and after a moment herself replies:

" 'Happily!'

"Reverently, for she is in the presence of the conscious and the benign—so it seems to her—she approaches the cord of corn and with her left hand takes of the selected ears along the top, an ear for each finger (that is, four), then with the right hand an equal number, placing them in the tray. She brings these forth and assisted by the male head of the household, shells them with such care that not a kernel is lost. Dust from the old bunch of soot is scattered over the shelled corn, and a curious sacred pigment is prepared, in an earthen ladle, of yellow paint and a kernel of salt, from the mountain near the lake of the dead, and the salt lake in the South. To these ingredients are added two or three kinds of little yellow flowers, the principal variety being precious in the eyes of the Zuni, as that which was left over of the seed stores of the gods. All this is mixed with pollen and water, and the whole tray of kernels is thoroughly sprinkled and anointed by stirring. The corn grains thus treated are bright-yellow in color and pleasantly odoriferous. All this is done that the 'seed' may have the power of reproduction, rapid growth and strength, and that it may bear fruit possessed of the properties of food, which fruit shall mature with the season when thrive most and bloom the little yellow flowers,—early autumn.

SUCCESSFUL SEED SELECTION.

1"Zuni Breadstuff," V, Frank H. Cushing,
The Millstone, Vol. IX, Nov. 5, 1884.

"We are at first surprised when we learn that to a remarkable degree the corn thus treated has vigor and the quality of ripening early; but our wonder may be lessened when we reflect that these seeds are the most perfect of the whole harvest, selected mostly from among those ears which soonest reach maturity. Still, with the Zuni all these things are living testaments of faith, proving the infallibility of his theory of Medicine or Fetichism and of his practice of religion."1

Mr. Cushing lived with the Zunis for a number of years and became a member of the tribe. He succeeded to a remarkable degree in attaining the Indian point of view, but appreciation of the Indians did not lessen his zeal for accuracy. This series of articles written in a charming literary style tells not only what the Zunis eat and how they secure their food, but gives a tantalizing glimpse of the character and personality of the Zuni.

In other articles of the same series the cultural methods of the Zunis are described. Though mixed with superstition, the methods employed are seen to be admirably adapted to supply fertility to the soil and to conserve moisture, enabling these Indians to produce maize under conditions that prohibit the growing of this crop by the methods ordinarily employed by whites.

2The description of the drought resisting adaptations of this type of maize has been published in The Journal of Agricultural Research, Vol. 1, No. 4, Dept. of Agriculture, Washington, D. C., January 10, 1914, where the subject is treated in a somewhat more botanical way,

The many incidental references to peculiar agricultural practices in Mr. Cushing's articles suggest that careful study of the agricultural practices of the different Indian tribes might disclose many facts of economic importance to agriculture. The results described in the present paper show one such fact: that we must thank the Indian for calling our attention to deep planting as a factor in drought resistance and for having developed a type of maize with peculiar characteristics that permit the utilization of this factor to an extraordinary extent.2

VALUABLE LORE NEGLECTED.

The development of maize as a cultivated plant must have involved a long series of unconscious experiments which resulted in important agricultural discoveries. That the results of these experiments are clothed in the language of myths and legends should not obscure the fact that in many instances they reveal sound agricultural principles. The study of this primitive agricultural lore has too long been left to the ethnologist. We have accepted the Indian's gift of maize but have hastened away without stopping to learn its full value or how best to utilize it.

Had we taken the trouble to learn and appreciate the Indian's discrimination in the choice of varieties, the differences in flavor and adaptability to different methods of cooking, we would not have assumed this valuable human food to be useful chiefly for the nourishment of our domestic animals.

Indians of the Southwest have preserved from pre-Columbian times a type of maize able to produce fair crops in regions where the better-known varieties of the East fail for lack of sufficient water An important factor in the drought resistance of this type of corn is its ability to force the growing shoot of the seedling to the surface of the soil when planted at a depth of a foot or more. At such depths less specialized varieties die before reaching the surface.

The literature of corn contains reports of many experiments conducted to determine the proper depth of planting, but the results are confusing and contradictory. It has generally been realized that the optimum depth is influenced by differences in soil and climate, but that the proper depth might vary with different varieties seems not to have been appreciated. The varying behavior of types when planted at different depths is additional proof that it is unsafe and unscientific to generalize with respect to cultural factors without taking type, varietal and even individual differences into account.

The primary root or radicle which is the first organ to emerge from the germinating seed is soon followed by the shoot or plumule. Enclosing the shoot is the cotyledonary sheath or coleoptyle, a tubular organ which is closed and pointed at the upper end. Between the base of the coleoptyle and the seed, the axis is somewhat elongated (see Fig. 5). With seeds germinated in the laboratory this elongation is so slight that it might easily be overlooked. Nevertheless, this small organ has not escaped the notice of morphologists and its nature has been the subject of much discussion. It has been called variously hypocotyl, mesocotyl and epicotyl. By some it is held to be an internode, by others merely an elongated node.

SUPERIORITY OF NAVAJO MAIZE FOR DEEP PLANTING
Test plantings of the Navajo dry-land maize, and Chinese and Boone County White, were made in a box at the depths shown in the diagram. The Navajo surpassed its competitors in growth at all depths; but from the lower levels it was the only one to emerge, due to its extraordinary adaptation for such growth, through the elongation of its mesocotyl. The diagram shows that at intermediate depths the Chinese and Boone varieties could not force their coleoptyles to the surface, and were obliged to make the last few centimeters of the distance by the aid of the true leaves, which in general are ill-adapted to pushing through solid earth. (Fig. 7.)

HOW THE SHOOT REACHES DAYLIGHT.

When a grain of corn germinates in the ground this usually insignificant organ is of vital importance to the life of the plant, for it is through the elongation of this mesocotyl that the shoot is enabled to reach the surface.

So long as a maize seedling remains below ground away from light, the mesocotyl will continue to elongate, until it reaches a maximum length that varies in different varieties of maize. As the mesocotyl elongates, the coleoptyle with its firm, sharp point is pushed upward through the soil. As soon as the coleoptyle emerges from the soil the elongation of the mesocotyl ceases and elongation of the internode bearing the first true leaf begins, forcing open the coleoptyle.

If the seed is planted so deep that the maximum elongation of the mesocotyl fails to bring the coleoptyle to the surface the task of penetrating the soil and reaching light devolves upon the first true leaves. In comparison with the sharp coleoptyle, these leaves are but poorly adapted for forcing their way through the soil, and if the tip of the coleoptyle stops more than a few centimeters below the surface these leaves usually crumple and never reach the light.

In the varieties of maize commonly grown the mesocotyl can seldom be forced to a length greater than four inches, while in the Hopi and Navajo varieties this usually minute organ may reach the relatively enormous length of 10 or even 12 inches, thus making it possible for these Indians to plant their maize deep in the ground where the soil is moist and germination is assured.

The mesocotyl is a beautiful contrivance for removing the young seedling from the seed and planting it at the proper depth from the surface. The true base of the plant is the base of the coleoptyle, the point from which the permanent roots arise. Since elongation of the mesocotyl ceases when the tip of the coleoptyle reaches the light, the length of the coleoptyle determines the depth at which the first permanent roots develop.

Figure 6 shows a seedling of Hopi maize as it developed when planted eight inches below the surface of the ground. The first permanent roots can be seen developing at the base of the coleoptyle 7 1/2 inches above the seed.

SPECIALIZATION IN HOPI SEEDLINGS
The varieties bred by the Indians of the southwest have but one seminal root, as shown in the three seedlings at the left (fig. 8); and they are able to throw their entire energy into the prolongation of this single radicle, so that they can force it down into moister ground, as the moisture recedes during periods of drought. Ordinary varieties like the Chinese, two seedlings of which are shown at the right (fig. 9). throw out several seminal roots and are obliged to divide their energy; thus they can not reach to the depths which the single radicle of the Hopi variety attains. In the struggle for existence during drought, the Hopi variety thus possesses a decided advantage.

GERMINATION OF NAVAJO MAIZE.

In the fall of 1912, W. T. Swingle and K. F. Kellerman of the Bureau of Plant Industry visited the region about Shiprock, N. M., in the Navajo reservation. and secured specimen ears of the maize grown by the Navajos. This collection was kindly placed at the disposal of the writer. Additional seed was later secured through the courtesy of Wm. T. Shelton, agent at Shiprock.

It has been frequently stated that the Navajos, like their neighbors the Hopi and Zunis, plant maize at unusual depths, 6, 12 and even 18 inches having been reported. Since planting at such depths was known to be impracticable with other varieties, an experiment was planned to test the ability of the Navajo maize in piercing the soil. A box, 70 cm. by 33 cm. and 34 cm. deep was sunk in the ground. A quantity of sandy loam soil sufficient to fill the box was slightly moistened and carefully sifted. At one end the box was filled to within one cm. of the top, the soil sloping in a straight line to within one cm. of the bottom at the other end.

Five seeds each of Navajo, Boone County White, and Chinese maize were placed in a row transverse to the inclined surface of the soil, two cm. from the top of the box.

A similar row was planted at four cm. from the top, and so on at the following depths: 6, 8, 10, 12, 16, 20, 24, 28 and 32 cm. The box was then filled with the soil and struck off level with the top. The seeds germinated promptly, and when the most advanced seedling had reached a total height of about 60 cm. the plants which appeared above the surface were dug up and the mesocotyl and coleoptyle were measured (see Fig. 7).

Twelve cm. was the greatest depth from which Chinese seedlings appeared at the surface. Boone seedlings appeared from all depths up to 20 cm. while Navajo plants appeared from all plantings to the very deepest, 32 cm.There were numerous instances where the combined length of the mesocotyl and coleoptyle was less than the depth at which the seed was planted. This, of course, means that the upper layers of the soil were penetrated by the true leaves. The maximum depth of soil thus penetrated by the true leaves of the plants of the Chinese variety was five cm. One Boone plant forced its leaves through eight cm. of soil. In all of the Navajo plants the coleoptyle reached the surface.

The extent to which the Chinese and Boone seedlings were able to penetrate the soil by means of the true leaves was doubtless much greater in the carefully prepared soil of the experiment than would be the case under field conditions where any slightly compacted lump of soil would deflect the tender leaves and cause them to crumple. On the other hand, many seedlings failed to come up where there was less than two cm. between the top of the coleoptyle and the surface of the ground. The results clearly show that the coleoptyle is the proper organ for penetrating the soil and where this office devolves upon the leaves, germination is uncertain.

It has been observed in many field plantings that the spatulate first leaf, formerly called the cotyledon, is the first evidence of the germinating plant. When this occurs in any considerable proportion of the plants, it is safe to assume that the seed has been planted too deep for the best results.

In examining these experimental plants it was observed that the root system of the Navajo variety differed from that of the other varieties. The roots of the Navajo seedlings extended to a greater depth, but there was only a single root arising from each seed, while in the Chinese and Boone seedlings the roots were shorter and more numerous. Further experiments with Hopi and Zuni varieties showed them to be like the Navajo variety in producing but a single root from the seed.

The roots of maize are of two kinds, those that rise from the embryo or seed, called seminal roots, and those produced from the nodes of the plant. Of the latter class, those that arise from the nodes above the ground are often called brace roots or aerial roots. In the varieties commonly grown in the United States there are, in addition to the primary root or radicle, from two to six additional roots that arise from the base of the cotyledon. These secondary seminal roots, though appearing somewhat later, usually equal or exceed the radicle in size. In the Pueblo varieties of maize these secondary seminal roots have been absent in all seedlings thus far examined, the radicle being the only root arising from the seed (see Figures 8 and 9).

ZUNI PLANTATION OF MAIZE IN ARIZONA
The soil is sandy and, except during rains, is quite dry on the surface. The seeds are planted at five to seven inches below ground, where there is sufficient moisture to germinate them; and as the ground dries out to lower levels, they thrust their single seminal roots farther down, so that they remain green and vigorous while plantings of ordinary varieties of maize under similar conditions dry up and perish. This photograph was made near Black Rock, Arizona. (Fig. 10.)

FIELD STUDIES OF PUEBLO MAIZE.

In September, 1913, opportunity was afforded for a short visit to the Zuni, Navajo and Hopi Indian reservations of Arizona and New Mexico. It was thus possible to form some idea of the agricultural significance of the peculiarities and habits of germination of this type of maize.

The value of deep planting made possible by the greatly elongated mesocotyl was obvious. In the localities selected by the Indians for planting maize the soil is sandy and in the absence of spring rains the surface layers are, of course, very dry. The seed, to germinate at all, must be planted deep enough to be in contact with the moist soil. In Navajo fields near Tohachi, N. M., plants were dug up and the remains of seeds were found at depths ranging from five to seven inches below the surface. Similar depths were found in a Zuni field near Black Rock, Arizona (see Figure 10.) In a Hopi field at Polacca, Arizona. near the first mesa, where the conditions are extreme, the seed had been planted at 10 inches from the surface. It thus appears that there is no fixed depth for planting, the custom being to plant deep enough to place the seed in moist soil. If the seed were planted at ordinary depths, germination might be delayed until the latter part of June or the first of July, at which time the rains usually occur, or if the seeds germinated as a result of one of the occasional showers occurring in May, the plants would die from subsequent dessication.

Like the long mesocotyl, the simple radicle of the Pueblo maize may be looked upon as an adaptation to the extreme conditions that exist where this type is grown. For six or eight weeks after planting no rain can reasonably be expected and during this time the moisture is constantly receding from the surface. By concentrating the energy of the seedling into a single root the latter is forced to greater depths and consequently kept in soil that is more moist than would be the case where a number of seminal roots developed.

Thus under ordinary conditions where moisture is distributed through the entire seed bed the seminal roots become of little importance as soon as the seedling is established and nodal roots have developed. If a half grown or nearly mature corn plant is carefully dug up, the seminal roots and traces of the seed can still be found but they are usually dry and shrunken and are obviously of little use to the plant. This was also the condition found in Navajo and Zuni maize fields, though the seminal roots were more strongly developed than in the eastern varieties. But, in the more extreme conditions existing in the fields near the Hopi villages, where, as stated, the seeds were planted deeper, it was found that the seminal roots were relatively much larger and were still alive and fresh, making it apparent that they retain their function of supplying moisture, and are able to play an important part during the entire life of the plant.

HOW THE HOPI INDIANS GROW MAIZE
Field at the base of the first Hopi mesa, near Polacca, Arizona. The hills are planted about 20 feet apart with 10 to 20 plants in a hill. The soil was apparently pure sand washed down from the mesa by winter rains, and was so dry that not a weed was to be found. In such unfavorable conditions the maize seeds planted at a depth of about 10 inches (25 cm.) had sent down their single, vigorous radicles or seminal roots to a depth of 14 or 15 inches more, so they were drawing moisture from a depth of at least two feet below the surface. No variety of corn cultivated in the eastern United States could possibly survive under such extreme conditions. (Fig. 11.)
MAIZE GROWING UNDER DIFFICULTIES
With most of the improved varieties of maize, the plant's chief concern seems to be to produce leaves and stalk. The ear is the last thing to receive its attention; in consequence, the ear is the first part of the plant to suffer under extreme drought or other unfavorable conditions. The habit of the Pueblo varieties is just the reverse; they seem determined to produce seed, even if they accomplish little else in life. A plant of this kind is shown here, where the single ear is more than one-half the height of the entire plant. (Fig. 12.)

HOW THE MAIZE IS GROWN.

NAVAJO MAIZE GROWN UNDER IRRIGATION
A single plant photographed, with the leaves and husks removed, at Shiprock, New Mexico. Although these Indian types of maize are particularly adapted to dry-farming, their yield under irrigation compares favorably with that of the ordinary commercial varieties. (Fig. 13.)
In one field at the base of the first Hopi Mesa the hills of maize were planted about 20 feet apart with from 10 to 20 plants in a hill (see Figure 11). The soil was apparently pure sand washed down by the winter rains and entirely destitute of vegetation other than the planted maize. An average hill dug up in the field was found to contain 15 plants ranging from 60 to 90 cm. in height. The remains of the seeds were found at 25 cm. from the surface and from each seed there descended a single large seminal root. These seminal roots were traced to a depth of 35 cm. and extended even farther down. They were still fresh and densely covered with fine branches. This mass of 15 seminal roots, while less in volume than the nodal roots arising near the surface, was apparently playing an important part in the support of the plants. The mesocotyls connecting the seminal roots with the plants above, while dry on the outside, were filled with live tissue quite unlike the dry and shrunken mesocotyls found in plants of similar age grown under more favorable conditions.

When planted by the Indian methods, the Hopi and Navajo varieties of maize have been found superior to the more improved eastern varieties for these very dry regions. At the time of our visit there was a small field near Keams Canyon that had been planted by eastern methods. The plants were in rows and thinned to one stalk to the hill. There had evidently been a fair germination but the plants had died without reaching maturity and had produced no seed. At the same time in the nearest Indian fields at Polacca the plants were dark green and maturing a fair crop, though the season was said to have been unusually dry.

Under extreme drought or other unfavorable conditions the persistent tendency of this type of maize to produce seed is very striking. Where any growth is possible it seems to be expended in the production of seed. Many plants were seen where the length of the ear was equal to one-half the height of the entire plant. A plant of this kind is shown in Figure 12. With most of the improved varieties of maize the ear is the first instead of the last part of the plant to suffer when unfavorable conditions are encountered.

Even under irrigation the somewhat larger strains grown by the Navajos have been found to compare very favorably with eastern types. Several acres of Navajo maize were seen at Shiprock, N. M., under irrigation. The fields were very uneven, apparently the result of alkali, but in the better portions the yield was good. The plants were standing about two feet apart in the row, the rows four feet apart, and nearly every plant was bearing from two to four fair sized ears (see Figure 13). The ears from 36 plants representing a number of distinct types were collected. The 36 plants bore in all 94 ears weighing 37.6 lbs., an average of 15.2 oz. per plant. The plants producing these ears averaged only a little over five feet in length.

CONCLUSIONS.

American agriculture is under obligation to the American Indian for having developed the maize plant to a high state of efficiency and for having adapted it to a wide range of environment.

See also, Collins: Tropical Varieties of Maize (1918)