ANCIENT SEEDS; SEED LONGEVITY

Association of Official Seed AnalystsSociety of Commercial Seed Technologists

ANCIENT SEEDS; SEED LONGEVITYAuthor(s): Vivian K. TooleSource: Journal of Seed Technology, Vol. 10, No. 1 (1986), pp. 1-23Published by: Association of Official Seed Analysts and the Society of Commercial Seed TechnologistsStable URL: http://www.jstor.org/stable/23433002 .

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ANCIENT SEEDS; SEED LONGEVITY

Vivian K. Toole1

Abstract

This paper tells about the widespread publicity, following archaeo

logical excavations, given to "mummy" grain and its viability, identifica tion of ancient grain as to species, findings of wild species, species grown in ancient times and conclusive evidence that mummy grain is nonviable. A brief reference is made to the longevity of the lotus seeds from Pulan

tien and to herbarium and buried seeds, and to present knowledge of

seed viability under controlled conditions, to imbibition, dormancy and

hard-seed effects on longevity and a glimpse into Alladin's lamp of seed

immortality.

Additional index words: buried seeds, mummified seeds, viability.

In the morning sow thy seed, and in the evening withhold not thy hand; for thou knowest not whether shall prosper, either this or that, or whether both shall be alike good. Ecclesiastes 11:6

Man has always sought eternal life; it is the theme in all religions. Since ancient times the seed has symbolized life and its renewal. We want to believe the stories of ancient seeds being viable but what about the facts? Do seeds have a secret of immortality which, if we could fath om it, would unleash the key to eternal youth?

ANCIENT SEED! Wheat has been found in the sarcophagi of ancient

Egyptian mummies. "The oldest tombs containing wheat belong to the

First Dynasty and are about 6,000 years old" (Buller, 1919). "Archaeolo

gists have discovered wheat in the rubbish heaps of the lake dwellings of both Switzerland and Italy, so that we have the clearest evidence that

this cereal was cultivated by prehistoric man. Unger found wheat in a

brick of the pyramid of Dashur in Egypt, to which he assigned the date 3359 B.C.; and the Chinese grew wheat as long ago as 2700 B.C. The ancient civilizations of Babylonia, Egypt, Crete, Greece, and Rome were

undoubtedly based on wheat as one of the principal food plants" (Buller, 1919).

Ancient records tell us that the introduction of agriculture opened the

way to civilization; settled communities based on agriculture were already established many thousands of years ago. Mr. Scott Elliott concluded that based on archaeological evidence, "the best guess as to the date of

W.S. Department of Agriculture, Seed Research Laboratory, Building 006, BARC

W, Beltsville, MD 20705.



2 JOURNAL OF SEED TECHNOLOGY

the first harvest is perhaps between 15,000 B.C. and 10,000 B.C." (Bnller, 1919). "The remains of neolithic man seem to prove that the growing of wheat was associated with his development." (Buller, 1919).

The wild form of wheat was first discovered by Körnicke in 1873

among herbarium specimens at the National Museum of Vienna (Buller,

1919). It had been collected in 1855 at Rasheyya, on the northwestern side of Mount Hermon in Palestine. Later, in 1906, it was found by Aaron Aaronsohn (Director of the Jewish Agricultural Experiment Station at

Haifa) growing near Mount Hermon. He collected many forms of this

plant; some resembled Triticum durum Desf. (durum wheat), others T. monococcum L. (einkorn).

The scientific name of the wild wheat used by Aaronsohn is T. dicoc cum dioccoides Aaronsohn because he thought it was most closely related

to emmer wheat, T. dicoccum Schrank. Aaronsohn established that the

wild wheat was indigenous to the region of Mount Hermon and the north

ern part of Trans-Jordan. His observations were confirmed by Cook of

the USDA. However, Cook felt the triple name made the wild wheat a

variety of the cultivated wheat. At his suggestion the name was changed

to T. hermonis Cook (quoted by Buller, 1919). The change in nomen clature gave the wild wheat the status of an independent species and commemorates Mount Hermon, its place of collection.

"The species of grain of the cultivation of which we have the oldest records is emmer. It is true that durum wheat has been found in Egypt

in some tombs of the first dynasty—that is four thousand years before the

Christian era—but emmer is found both in far greater abundance and in

all of the tombs. It is not at the present time cultivated anywhere in

Egypt, durum wheat having since historic times taken its place. Emmer

has been found in the lake dwellings of Wangen and Robenhausen, which date back to the end of the neolithic epoch, a little before the bronze

age. This, therefore, is the only species which has been cultivated from the very beginning of civilization, and we are justified in asserting it to be the progenitor of our cultivated wheats" (Aaronsohn quoted by Buller,

1919). Percival (1921) also refers to emmer as being "widely grown by prehistoric man in various parts of Europe and Asia, and was one of the

chief cereals cultivated in Egypt from the earliest times to the Graeco Roman period and later."

"Barley is stated to be one of the first cereals cultivated by man.

Grains of barley have been discovered in Egypt belonging to pre-dynastic

and early dynastic periods" (Luthra, 1936). The samples found at Mohen

jodaro are believed to have remained buried for about 4,000 years and

are of a later date than the Egyptian mummy wheat.



VOLUME 10, NUMBER 1, 1986 3

Aberg (1950) identified, by taxonomic characters, 5,000 year old barley and wheat samples from the Saqqara Pyramid in Egypt as closely related to Hordeum vulgare L. emend Lam. var. pallidum, Ser. and two species of wheat as T. monococcum L. and T. dicoccum Schübl. of the emmer

type. The seeds were brownish in color because of ageing. The Saqqara barley had several morphological characters similar to barley from the neolithic period excavated in the northern Fayum desert and is closely related to the Manchuria-type of today (Manchuria was derived from

Manshury, village in the Nile). According to Aberg, the ancient Egyptians knew abut T. monococcum and even cultivated it. The emmer type

closely resembled the emmer type of wheat from 1400 B.C., illustrated

by Percival (1921).

VIABLE ANCIENT SEED! Words to capture the imagination of man in his eternal search for earthly immortality—Ponce de Leon's Fountain of Youth.

In spite of research, popular belief clings tenaciously to the tales con

cerning the germination of seeds from ancient tombs. We read in the

Gardner's Chronicle of 1843 (Anon, 1843) of seeds possessing the power "under certain circumstances, of preserving their viability for an appar ently indefinite period." In this same Chronicle they give the following

story which appeared in the London Times of September 1840.

"Sir Gardiner Wilkinson, when in the Thebaid, opened an ancient tomb (which had probably remained unvisited by man during the greater part of 3000 years), and from some alabaster sepulchral vases therein took

with his own hands a quantity of Wheat and Barley that had been there

preserved. Portions of this grain Sir G. Wilkinson had given to Mr. Petti

grew, who presented Mr. Tupper with 12 grains of the venerable harvest." In 1840 Mr. Tupper "a most exact and conscientious man," sowed these

12 grains, care being taken to insure that no other grain was present; one

germinated. The nourished seedling produced two ears and says Mr.

Tupper—"If, and I see no reason to disbelieve it, if this plant of Wheat be indeed the product of a grain preserved since the time of the Pharoahs, we moderns may, within a little year, eat bread made of Corn which

Joseph might have reasonably thought to store in his granaries, and almost

literally snatch a meal from the kneading-troughs of departing Israel."

(Anon, 1843).

Then there's the story about "an Englishman of unimpeachable hon

esty" who "brought home from Egypt some seeds which were undoubted

ly genuine; it was quite certain that they were several thousand years

old. The gentleman had these seeds planted in his garden and to the amazement of 'those in the know,' they produced a very fine crop, in




distinguishable from modern varieties. On further investigation, however, it was found that the gentleman's gardener, thinking it most unlikely that the poor-looking seeds his master had brought home from Egypt would

produce much of a show, had planted modern seeds alongside the old, in order, he said, not to disappoint his master" (Anon, 1951b).

According to Mr. Grimstone, "Mummy" pea was found in a hermetic

ally sealed antique vase in a mummy pit in Egypt and was computed to

have lain there about 3,000 years (Anon, 1849). The accounts vary (Anon, 1847; Anon, 1873; Lane 1867). In some cases Egyptian pea seeds were

found "in the hand of a mummy of a young girl which was excavated

from a tomb nearly 5,000 years old. All the seeds germinated and when

the plants matured blue morning-glories (or sweet peas) 'looking like a

tiny Egyptian face' were produced" (Anon, 1922). Scientists at the Brandy

Experimental Farm, University of Virginia were amazed at an article en

titled "Peas from King Tut's tomb flourish in Florida" that appeared in

1945 (Anon, 1945). They responded with a disclaimer article (White,

1946).

"This question of viability was brought forward again during the dis

covery and examination of the tomb of King Tutankhamen in 1923, by Mr. Howard Carter and Lord Carnarvon, and has received attention from

various quarters since that date. Now another claim has been made by

an American farmer that wheat taken from the tomb of Tutankhamen has

been made to grow, and this fact has received much publicity in the press"

(Anon, 1931). As late as 1951, in spite of many negative results on ger mination of "mummy" grain, publicity was again given to viable grains from the tomb of Tutankhamen in a Washington, D.C. newspaper (Justice and Bass, 1978).

I could quote many such stories, but let us turn from the sensational

news and look at the facts. Tombs in modern times have been used for

storage of grain, mummies have been packed for shipment in imperfectly threshed straw and the combination of a clever guide and the credulous tourists—each could contribute to the popular tales.

Bower (1923) in his Botany of the Living Plant mentions that "A. de

Candolle, after examining the evidence up to 1882, concluded that no

grain taken from an ancient Egyptian sarcophagus and sown by horti culturists has ever been known to germinate; nor is there any trustworthy

evidence up to the present date." Buller (1919) in his Essays on Wheat

says "It is still currently reported that this mummy wheat, after being sown, has been observed to germinate; but there is no truth whatever to

this story. Careful experiment has demonstrated that all real mummy wheat has entirely lost its vitality."




Another report is attributed to Sir John Marshall (quoted by Luthra,

1936) on some samples of cereals found at Mohenjodaro and kindly sup plied by the Director-General of Archaeology in India, lie says "The

grains in all the three samples are completely carbonized. They have

turned black both on the surface as well as inside and have the appear ance of charred material. The surface is quite smooth as in fresh normal

grains and both the proximal and distal ends are intact. The form and outline of the grain is very well preserved and they still possess their

typical shape. The embryo retains its form in some grains but in others it is disintegrated and a hollow is left. The grains were tested for ger mination power but were not found to be viable at all. On being mois tened with water the grains crumbled into powder forming fine black ash"

(Luthra, 1936). Identifications of such intact carbonized grain were given in the first part of this paper.

Barley seeds from the tomb of King Tutankhamen (c. 1350 B.C.) sent to the Cereals Research Station at St. Albans by the Chief botanist of the

Ministry of Agriculture, Egypt, also were extensively carbonized; their structure was undamaged although the weight and density was less than

that of fresh English barley (Barton-Wright et al., 1944).

John Percival (1921) of the Department of Agricultural Botany at

Reading University, provides further evidence. He states, "In many cases

they are completely carbonized, and in this state are merely friable masses

of charcoal with the form of the grain: such are those obtained from the neolithic pile dwellings of Switzerland and the pre-Roman settlements of this country. Specimens from vases deposited in the tombs of ancient

Egypt, where they have been kept dry, away from atmospheric changes, show much less alteration and, except that they are a dark reddish-brown

colour as if scorched, appear normal in size and shape." Others (Anon,

1922) report that microscopic examination of the embryo of wheat taken

from a vase in an ancient tomb was a brownish color and practically de

stroyed. Percival (1921) examined grain found by Professor Flinders Petrie in the Graeco-Roman Cemetery at Hawara (about first century

B.C.) and reported "These were plump, and similar to some forms of T. durum Desf. grown at the present day. The pericarp was a dark reddish brown tint, its structure unchanged. The aleuron layer had retained its

original shape, with normal cell walls and typical round aleuron grains within." Examples of carbonized and discolored seeds are given in Fig ures 1 and 2.

Gain (1900) also reports that wheat and barley probably 4,000 to 6,000 years old, from ancient tombs, are a reddish-brown color and although

they have an exterior appearance of good preservation, the embryo has

undergone a marked chemical change and is no longer viable.



JOURNAL OF SEED TECHNOLOGY

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Figure 1. Upper: Left to right: Ancient and Modern Wheat. Lower: Left to right: Ancient and Modern Barley. Ancient Carbonized Mesopotamium seed, 3500 B.C. from the Oxford field expedition, 1926, Prof. S. Langdon. Loaned for photograph ing by the Federal Seed Laboratory, Seed Branch, Livestock and Seed Division, AMS, USDA. Photograph by John W. Mitchell.

Grain from a model granery, found in a 19th dynasty tomb (ca 1200

B.C.) in western Thebes and brought to England by Sir Wallis Budge of the British Museum was tested for viability soon after by the National

Institute of Agricultural Botany at Cambridge (Anon, 1934; Brooke, 1935)



VOLUME 10, NUMBER 1, 1986

Fa

Figure 2. Left: Ancient Egyptian Wheat (Emmer), found in two graves of the "mid dle kingdom" (about 1900 B.C.) in the neighborhood of the pyramid temple of

King Ne-user-re (2600 B.C.) at Abusir, near Cairo, Egypt (Gift of Deutsche Orient-Gesellschaft (1903). Loaned for photographing by the Federal Seed Lab., Seed Branch, Livestocks and Seed Division, AMS, USDA. Right: Modern Wheat. USDA Photographer.

and by the Dominion Seed Branch of Winnipeg (Brooke, 1935) and at

Kew (Anon, 1931; Anon, 1933). These tests showed that the wheat had

entirely lost its viability; all seeds decayed.

Maize grains, from an archaeological site in northeast Arizona around

600 years old, failed to germinate under conditions that caused modern

maize to germinate (Derbyshire et al., 1977). Light microscopy and elec tron microscopy revealed intact endosperm tissue similar to that of mod

ern maize but the embryos were discolored and damaged. Protein was

found in both the endosperm and embryos of the aged seeds but not in the

same proportion as in modern maize.

The Agricultural Research Council's Unit of Developmental Botany,

Cambridge (Osborne, 1974), provides the most classical results on their

biochemical studies of ancient seeds. They used grains of different ages ranging from the last season's crop to one 6,000 years old. The oldest

grains tested were those of wheat from Egyptian tombs and from silos

of Fayum and are carbon dated 4,000 and 6,000 years respectively. They found that RNAase activity is present in wheat up to 100 years old, includ

ing a seed lot (1871 harvest) recently discovered in a-sealed bottle during demolition of a building in Kent. The two oldest lots showed no activity in any enzyme assay carried out. Their data on isolated embryos from

rye seeds of 9 to 15 percent moisture stored for six years showed that the

greatest impairment of protein synthetic activity occurs as the viablity

falls from about 95 to 85 percent. Electron microscope studies revealed




that neither tRNA nor DNA could be separated from the archaeological grains, indicating complete degradation of macromolecular nucleic acid constituents of the cells. The polysaccharide macromolecules were highly

resistant to degradation as indicated by structurally intact cell walls and

starch grains even in the oldest specimens. This is doubtless the reason

why non-viable seeds 6,000 years old can still be identified as to species. Thus no credence is attached to the sensational assertions of wheat re

maining viable for thousands of years.

Germination of 12, 22 and 0 percent is reported for seeds of barley, oat, and wheat, respectively, of about 8 percent moisture content which

were found in sealed glass tubes within the foundation stone of the

Nuremburg City Theatre after 123 years (Aufhammer and Simon, 1957,

quoted by Roberts, 1972a). Roberts (1972a) estimated the 'effective'

temperature during storage to have been around 10 C. The longest record

of viability (16 percent) for wheat is 25 years (Percival, 1921). This seed

from the Rothamsted farm was artificially dried after harvest and stored

in glass bottles.

Authentic records of seeds surviving for over a century come from

germination tests made on herbarium and museum samples of the Legu

minosae family known for producing hard seeds: Albizzia julibrissin Durazz., 147 years (Anon, 1942a, b); Cassia hicapsularis L., 115 years

(Recquerel, 1907, quoted by Youngman, 1951), C. multijuga (L.) R. Rich, 158 years (Becquerel, 1907, quoted by Youngman, 1951); Goodia latifolia Salisbury, 105 years (Ewart, 1908, quoted by Turner, 1933); Hovea het

erophylla A. Cunningham ex Hooker f., 105 years (Ewart, 1908, quoted by Turner, 1933); and Nehimbium 150 and 250 years (Robert Brown

quoted by Anon, 1942b; Thieret, 1954).

Records of survival in soil over 500 years are for seeds of Nelumbo,

Lupinum, Canna, Chenopodium and Spergula. In 1923, a Japanese botan

ist, Ichiro Ohga (1923, 1926a, 1927; Exell, 1931), discovered a layer of seeds of Indian lotus, Nelumbo nucífera Gaerth. (Nelumbium speciosum Willd.) in an ancient lakebed deposit at Pulantien in northeast China. After scarification, all 35 seeds tested germinated and the 15 seeds sent to other persons also germinated. Seeds from the same source also were

scarified and tested for viability in 1939 by Chamberlain of the Univer

sity of Chicago (Anon, 1939), and in 1951 by Wester of the National Park Service and Chaney of the University of California (Chaney, 1951b); all seed germinated. Lipids of unimbibed Pulantien seeds were examined

by Priestly and Posthumum (1982) and found to be still highly polyun saturated.

14Carbon determinations on the age of these lotus seeds are not in

agreement (Anon, 1951a; Godwin and Willis, 1964; Libby, 1954). Chaney




(1951a) indicates that these Pulantien fruits may have retained their via

bility over a period exceeding 400 years as was predicted by Ohga (1927). Wester (1973), on the basis of historical evidence he provides and its ex cellent agreement with Libby's "carbon determination of 1040 ± 200

years, states "it seems reasonable to conclude that the maximum age of

the seeds in the Pulantien deposit can be calculated from 371 A.D. which in 1951 would then recognize 1580 years as the maximum age of these

seeds." However, he stresses the need for further 1 'carbon dating on the

seed coats and embryos separately, as any error in radiocarbon dating

arises from the living embryo and not from the nonliving seed coat. Ohga (1926b) reports that the ancient lotus fruits were smaller and differed in

size, color, shape, luster and texture from recent fruits.

Radiocarbon dating for Lupinus arcticus S. Wats, seeds found in the Yukon in a rodent burrow 10-20 feet below the surface and for Carina seeds enclosed in a Juglans australis Griseb nutshell forming part of a rat tle necklace found in a tomb in northwest Argentina were based on a

rodent skull for the former and for the latter a camelloid bone from a

rubbish heap said to be of similar age to the tomb (Anon, 1968; Godwin, 1968; Porsild et al., 1967; Sivori et al., 1968): "In all such situations one lacks unequivocal evidence of the contemporaneity of the seed and the

material taken for radiocarbon or other dating. Such evidence leaves one

in doubt of the very ancient age attributed to Lupinus of 10,000 years and to Canna of 530 years" (Godwin, 1968). However, Lerman and Cig liano (1971) concluded that the age of the Canna was about 600 years old based on their new carbon-14 dates of the nutshell and a trash heap.

Odum (1965) in his survey of archaeological sites in Denmark and Swe den reported viability of 100 and 600 years for some weed seeds and about

1700 for seed of Chenopoclium album L. and Spergula arvensis L. Archae

ological dating was criticized as being 'indirect and weak'; dating should be on the seeds themselves (Ewart quoted by Bewley and Black, 1982,

and Bewley and Black, 1982).

The most accurate method of determining longevity in nature is ob

viously to germinate seeds that have been held in the soil for a known

period of time. In 1879, Beal (1905) prepared 20 sets of bottles each with 50 seeds of each of 22 different species of plants mixed with sand. The

inverted, narrow-mouthed, unstoppered bottles were buried in the soil

at a depth of 18 inches, each inclined at an angle to prevent water accu

mulation. Tests were made at 5 year intervals through the 40th year and

then at 10 year intervals. When tested at the 80th year of burial (Darling ton and Steinbauer, 1961) the only survivors were Oenothera biennis L.

(10%), Rumex crispus L. (12%), and Verbascum blattaria L. (70%). After 90 years (Kivilaan & Bandurski, 1973) only Verbascum survived




with 20% germination. After 100 years (Kivilaan & Bandurski, 1981) V. blattaria germinated 42% and V. thapsus L. and Malva 2% each. Malva had not germinated since the 20th year and V. thapsus since the 35th year.

In 1905, Duvel buried seeds under more natural soil conditions. The seeds were mixed with sterilized soil in an earthenware pot and covered

with a porous earthen saucer. Each pot contained 100 or 200 seeds of a

given species except in the case of the larger seeds fewer numbers were

used. In all, 33 sets of pots were buried at a depth of 8, 22 and 42 inches. Each set represented 112 samples of seeds and 107 species. Tests were made at various intervals. The general tendency was for seeds buried at

the 8 inch depth to germinate less than those from the 42 inch depth, be cause of the more equable conditions of the latter. After 39 years (Toole and Brown, 1946) in the soil at 22 and 42 inch depth, 16 species, repre senting 10 plant families, germinated above 15% from at least one depth: Abutilón theophrasti Medik, Ambrosia artemisiifolia L., Convolvulus sep ium L., Datura stramonium L., Ipomoea lacunosa L., Lespedeza interme

dia (S. Wats.) Britton, Nicotiana tabacum L., Oenothera biennis L., Ono

pordum acanthium L., Phytolacca americana L., Potentilla norvegica L., Robinia pseudoacacia L., Rudbeckia hirta L., Solanum nigrum L., Trifo lium pratense L., and Verbascum thapsus L. The high germination for Datura (91-88%), Phytolacca (81-90%) and Solanum (83-79%) indicates the possibility that these species might persist in the soil longer than the records for dry seeds in storage. Twenty other species showed some

life after 39 years albeit most were 6% viable or less. Three crop plants

showed some germination: tobacco (22-17%) red clover (2-3%) and Ken

tucky bluegrass (1-2%), however, 15 species of crop plants did not survive the first year. Unfortunately, regrading of the burial area by the United

States War Department in 1941 terminated the experiment. Seeds buried

in the soil lose viability fastest with shallow placement of seeds (Toole and Brown, 1946; Taylorson, 1970) and when they have little initial dor

mancy (Taylorson, 1970).

Blakeslee's (1954, and Avery et al., 1959) mutation studies with these Datura seeds buried 39 years showed only 5.1% segregation due to gene mutation. He concluded that the high mutation rate attained from seed

aged in his laboratory was probably the result of factors other than age alone. However, effect of age on mutation is recorded for many species.

Lewis (1973) placed seeds of 7 cereals, 15 grasses, 8 herbage legumes and 9 arable weeds in woven glass-cloth bags inside earthenware pots. The pots were placed in trenches and covered 13, 26 and 39 cm deep in three soil types. After 20 years, Ranunculus repens L. (53%) Chenopdium album L. (23%) and Rumex crispus L. (18%) showed the highest germin




ation. Phleum spp. was the only grass to persist for 20 years and most

legumes showed some germination.

A review by Crocker (1938) covers longevity of seeds recovered from soils of unknown age, seeds under water, and seeds that do not withstand

drying that are not covered here.

In the first half of this century, controlled seed storage received much attention. The wealth of information on this subject is covered in many reviews but only the most recent ones are listed (Bass, 1979, 1980; Justice and Bass, 1978; Roberts, 1972a; Roos, In Press). Briefly, with the excep tion of seed kinds that do not withstand drying, viability is best maintained at low temperature, low humidity (low seed moisture) and low oxygen pressure. In such a controlled seed storage study, it was observed that im

portant genes were lost in the continued growing of a crop. For example,

field plantings of Maryland Mammoth soybean seeds, after storage for 10

years under conditions that showed no loss in germination, were made in

comparison with those of seeds grown the previous year (Toole and Toole,

1946). There was no apparent difference in vigor of growth or in plant de

velopment except that plants from the new seed of Maryland Mammoth

had a more upright growth than those from the stored seed. The most

striking loss of vigor without loss of viability was reduction in size of all

seedling parts observed in snap bean seeds stored 47 months at 19 °C and 57 percent relative humidity (Toole and Toole, 1953; Toole et al., 1957).

Knowledge gained from such controlled environmental studies led to

the feasibility of maintaining for longer periods valuable germplasm for

breeding purposes. In 1956, Congress appropriated $450,000 for the con struction of the National Seed Storage Laboratory (NSSL) in Fort Collins, Colorado plus $100,000 per year to operate it (Bass, 1981). Such facilities are also found in Europe and Japan (Hawkes and Lange, 1973). The

NSSL's two-fold purpose is to preserve for posterity valuable genetic

characters of the original wild types and to research ways to keep seeds

vigorously viable for much longer periods.

The life of a seed varies greatly between plant families, within a genus and even cultivars of a species. The seed has its built-in mechanisms for survival: dormancy, hard-seediness, impermeability to water and

gases, the light mechanism (phytochrome), hormonal control (promoters and inhibitors), etc. However, from seed formation until death, environ mental factors exert an influence over the genetic make-up of a seed and

its longevity. The seasonal periodicity of dormancy of seeds buried in the soil changes the sensitivity of the seeds to their environment and is of high survival value (Baskin and Baskin, 1977, 1981; Karssen, 1982, Roberts, 1972b; Roberts and Neilson, 1982; Wesson and Wareing, 1969). Outside

agents (microflora, insects, ionizing radiation, etc.) can also exert an in




fluence on the seed's longevity in air-dry storage (Christensen, 1973; Howe, 1973). The weak and mechanically damaged ones die first and the

vigorous ones live for a longer period.

In the case of Nelumbium seeds, Ohga (1926a and b) tells us that its fruit coat, which encloses the seed, is impermeable to water and gases.

Although those seeds listed as retaining viability for over a century (her barium, museum) were of the hard-seeded type, many of the species re

ported above as viable after 39 years burial are not hard-seeded but had absorbed moisture while in the soil.

Lettuce seeds held imbibed while stored at a high temperature and

high humidity become dormant and remain viable but air-dry seeds held under the same conditions lose viability rapidly (Toole and Toole, 1953). This has been verified for lettuce (Lactuca sativa L.) and shown for ash

(Fraxinus americana L.) (Villiers, 1972). Why is it that fully imbibed seeds under a non-germinating environment have a greatly extended life

span while an increase in seed moisture content of seed stored air-dry

reduces the longevity? The following explanation is provided by more recent research (Villiers, 1975, 1980). Although the metabolic rate is not

high in imbibed dormant seeds, they are capable of undergoing enzymatic reactions and of manipulating large structural units such as proteins or

nucleic acids. Repair enzymes cannot function in the absence of suf

ficient water content in air-dry seeds. The concept that the viability of

normal tissues in imbibed seeds under a non-germinating environment

(induced dormancy) is maintained structurally and functionally by repair

mechanisms or are replaced as whole units by new organelles sheds new

light on the maintenance of viability in imbibed lettuce and other seeds.

In air-dry storage the environmental conditions of storage rather than

the length of storage is the important factor in maintaining viability. The

physiological and biochemical changes taking place in storage are a func tion of conditions of storage: induction and loss of dormancy, shifts in

temperature requirements for germination or for maintaining dormancy,

occurrence of seedling abnormalities, loss of vigor, loss or reduction of

enzymes, loss of membrane integrity, auto-oxidation of lipids, etc.

Almost every system within the seed is involved in the series of

changes leading to loss of viability. Roberts (1972c) classifies the theories on loss of viability as extrinsic or intrinsic (Figure 3). Extrinsic theories

suppose that loss of viability is due to outside agents (previously discussed) and intrinsic theories suggest that deterioration is a result of the inherent

nature of seeds and their molecular changes. Several reviews cover de

terioration due to intrinsic changes (Anderson, 1973; Anderson and Baker,

1983; Bewley and Black, 1982; Cheng, 1984; Cherry, 1983; Ilallam, 1972; Halloin, 1983; Harman, 1983; Harrington, 1973; Koostra, 1973; Mills, 1983;




Ionising j> radiations

Estrinsic ^

\f———/ Fungi

Pathogen attack

Mycotoxins

Growth inhibitors

Mutagens

Phenolics

Indoleacetic acid

Abscisic acid

/

Mitochondria

Plastids, / Denatu ration / lysosomes, of lipoprotein

\ dictyosomes, membranes

\ etc.

Cell membranes

Respiratory loss of food

Carbohydrates

Lipids

Loss of vilamins. hormones, etc.

Figure 3 from E. H. Roberts (1972c). A Classification of viability theories. Viability of Seeds. Syracuse University Press. USA, p. 283 (Table 9.5).

Osborne, 1980, 1982; Roberts, 1972a, b, c, 1973a, b, 1979; Roberts et al., 1967; Roberts and Ellis, 1982; Roos, 1982; St. Angelo and Ory, 1983).

Maintenance of the genetic integrity of seed populations both during storage and regeneration is of chief importance. During prolonged stor




age and in crop regeneration as in seed banks, genetic changes such as chromosomal aberrations and differential survival of genetically hetero

geneous accessions becomes increasingly important (Roos, 1982). The les son learned from the southern corn leaf blight in 1970 is that genetic uniformity limits the range of host/pathogen response and is the basis of

wide-spread vulnerability to epidemics. The uniform produce demanded

by the market made possible the feeding of an expanding population. However, most of our major crops with their highly increased production

are impressively uniform genetically and impressively vulnerable to epi demics (National Academy of Science, USDA, 1972). It is imperative that we preserve specific genes and genetic diversity (National Academy of Science, USDA, 1978).

Attempts to prolong seed storage life without genetic changes have

recently centered on the use of liquid nitrogen (LN2) as a storage medium

(Stanwood and Bass, 1978, 1981). At —196 C it is presumed that all bio chemical activity is reduced to essentially zero. Storage in vacuum (Went, 1969) and air-drying followed by freeze-drying to reduce seed-moisture to very low levels (Woodstock et al., 1983) are also used. They (see Hallam, Osborne, Roberts and Roos references) are probing the cause of

lesions that lead to senescence and death of the embryo and loss of their

repair mechanisms. The major lesions that are recognized and continue

to be studied involve membranes, mitochondria, ribosomes, post-ribosomal

supernatant components and the nuclei. Evidence does not indicate a

single underlying cause of deterioration. Another recent approach is to

use non-aqueous solvents such as acetone to impregnate seeds with chemi

cals to protect essential seed constituents such as RNA, DNA, proteins

and lipids from harmful oxidative reactions during storage (Woodstock et al., 1983).

In summary, archaeological findings of ancient seeds tell us much about ancient culture. Wheat is and has been of vast importance to the

world—the chief basis of ancient civilizations. Electron microscope studies revealed complete degradation of macromolecular nucleic acid constitu

ents. If fragments of the genetic system were still present in these an cient seeds, the genetic information could be identified and compared to that of present day cultivars by recombinant DNA and monoclonal anti

body techniques. Measurable amounts of DNA (Rogers and Bendich, In Press) have recently been extracted from mummified seeds of Encelia

virginensis (A. Neis.) Blake, Eschscholzia minutiflora S. Wats., Lycium shockleyi, Gray, Juniperus osteosperma (Torr.) Little, Opuntia ramosis sima Engelm. and Symphoricarpos sp. obtained from packrat middens and radiocarbon dated as 500 to greater than 44,600 years old. Although there was a general trend toward DNA degradation with age, some of




the samples contained high molecular weight DNA thus the ability to clone and sequence DNA from this material is a very real possibility. The

prospect of discovering more ancient seeds and decoding their genetic

makeup using this technique should be zealously pursued.

Ancient seeds are incapable of germinating; they are completely car bonized or of a dark reddish brown color. On the other hand, the intact structure of the grain due to the preservation of macromolecular poly

saccharides made possible the identifications of the seeds as to species; we know the kinds of cereals cultivated by ancient man. Also the pro

genitor of ancient cereals has been found in Palestine.

Nelumbium seeds appear to be one of the longest lived which is is attributed to water and gas impermeability of the fruit coat enclosing the seeds.

Burial in soil is better than air-dry storage for long-term preservation of seeds. Under environmental controlled storage, relative high humidity and high temperature causes rapid deterioration of air-dry seeds yet fully imbibed seeds under the same conditions become dormant and have an

extended life. The cellular repair mechanism maintains molecular or

ganization in the living seed; molecular disorganization is the state of the non-living seed.

We are still searching for ways to promote eternal life in seeds. The interest today is in long-term preservation of vigorous genetic materials:

air drying followed by freeze drying to reduce seed moisture, storage in

liquid nitrogen, invigorating treatments (as with PEG), nature and cause

of membrane damage, cellular repair mechanisms, etc. Perhaps the bio

chemist-physiologist should look at situations where the deterioration

processes move slowly as in the bean seeds that lost vigor but not viability.

The search for eternal youth continues but the battlefield is now in

the research laboratory rather than in the pyramids. The seed, containing a complete new individual, is indeed a provocative challenge in the study of senescence and longevity.

For this is the source of the root and the bud . . .

World unto world unto world remolded.

This is the seed, compact of God,

Wherein all mystery is enfolded.

"On a Seed" by Georgie Starbuck Galbraith.




Acknowledgment

I thank Mr. Cornelius L. McKissick, librarian, National Agricultural Library, U.S. Department of Agriculture, and Miss Sonya Whitted, Bio

logical Aid, Seed Research Laboratory, ARS, U.S. Department of Agri culture, for their library assistance and to Dr. Patricia Rhodes, Research

Plant Physiologist, Plant Molecular Genetics Laboratory, ARS, U.S. De

partment of Agriculture for suggestions on DNA.

Addendum

Recent archaeological investigations at Metaponta, Italy, by Dr. Jo

seph C. Carter and his associates, Drs. Lorenzo Costantini and Franco

Rollo, open the exciting possibility of decoding the genetic makeup of

ancient seeds by cloning and sequencing of DNA.

The first important discovery of ancient seeds by Dr. Carter, Director

of the Institute of Classical Archaeology at the University of Texas at

Austin, was in the 1978 archaeological excavation of a submerged sacred

spring and resevoir at Pizzica Pantanello in the rural territory of Metapon

to, a Greek colonial city in southern Italy. The remarkable state of pres

ervation of the seeds was attributed to the "anaerobic conditions created

by the ground water, flowing as it had in ancient times from the mouth

of the spring and into the reservoir." A pumping system used in the 1981 1982 studies made it possible to carry out a clean and precise excavation

of this area. (Carter, J. 1983. "Excavation in the sanctuary area, Pizzica

Pantanello, 1982" and Costantini, Lorenzo 1983. "Bioarchaeological Re

search at Pizzica Pantanello." In: The Territory of Metaponto 1981-1982.

Austin, Texas.

Seeds were found in the reservoir dated to the mid-fourth and early

third centuries B.C. Ranunculus (buttercup) seeds found in the mid

fourth century level indicated a humid but not thoroughly submerged ambience whereas the presence of Ceratophyllum (coontail) and Zan nichellia (horned pond weed) indicated a swampy condition, which cre ated a very compact stratum of organic material, in the level of the im

mediately succeeding period. The entire deposit covers the brief span of a half century. Seeds of Ficus carica L., (fig), Olea europaea L., (olive), Vitis vinifera L., (grape), Rubus sp. (blackberry), many weed and other seeds were found in both these levels. Seeds from the swampy level were

remarkably well-preserved. Carbonized Triticum dicoccum Schrank,

(emmer) and Triticum compactum Host, (wheat) and Hordeum vulgare L., (barley) seeds and weed, forage and fruit seeds were found in the levels dated mid-fourth century B.C. and in the nonswampy level imme

diately over the compacted organic material of the early third century

B.C. The absolute dates were established by a study of associated ceramic

and other archaeological materials at the site.




Seeds of Ceratophijllum, Zostera and more recently Vitis were carbon

dated by Salvatore Valastro, Radio Carbon Laboratory, Balcones Research

Center, Austin, Texas (Personal communication). These dates are in full

agreement with those determined from the archaeological context (Red

figured pottery).

Dr. Rollo (Personal communication) found clonable fragments of DNA in some of these well-preserved seeds and also in cress seeds (Lepidium

sativum L.) from Thebes dated approximately 1400 years B.C. (F. Rollo. 1985. "Characterization by molecular hybridization of RNA fragments isolated from ancient [1400 B.C.] seeds." Theor. Appl. Genet. 71:330-333.) He is presently trying to establish whether the DNA is endogenous to the

seed genome or is due to possible contamination by molds. If mold con tamination is excluded by these painstaking experiments he will then characterize and sequence the DNA. We anticipate that Dr. Rollo may

be able to distinguish fragments of coding from non-coding sequences and provide useful new information about the genetic composition of

ancient seeds.

Secondly, the remarkable preservation of the seeds from the swampy

area may support Dr. Villier's theory (see page 12 of this paper) that im

bibed seeds under a non-germinating environment (induced dormancy) maintain viability better than dry seeds because moisture is needed for

repair or replacement mechanisms.

Thirdly, Dr. Carter et al. state that the bulk of material relating to their study of the history of the rise and fall of agriculture at Metaponto during the third and fourth centuries B.C. was provided by the ancient

seeds and pollen from this excavation. (Carter, J. C., L. Costantini, C.

D'Annibale, J. R. Jones, R. L. Folk, and D. Sullivan, with the assistance of M. E. Reed. 1985. "Population and Agriculture: Magna Grecia in the fourth century B.C." In: Caroline Malone and Simon Stoddart eds.

Italian Archaeology IV, The Cambridge Conference. Part 1, The Human

Landscape. BAR International Series 243, Oxford, England. 1985.)

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Documents

ANCIENT SEEDS; SEED LONGEVITY