Courtesy of Herb.com | Robert Connell Clarke
GENETICS AND SEX IN CANNABIS
Sex is an inherited trait in Cannabis, and can be explained in much the same terms as human sexuality can. Like a human being, Cannabis is a diploid organism: its chromosomes come in pairs. Chromosomes are microscopic structures within the cells on which the genes are aligned. Cannabis has 10 pairs of chromosomes (n=10), for a total of 20 chromosomes (2m=20).
One pair of chromosomes carries the primary genes that determine sex. These chromosomes are labelled either X or Y. Male plants have an XY pair of sex chromosomes. Females have XX. Each parent contribute one set of 10 chromosomes, which includes one sex chromosome, to the embryo. The sex chromosome carried by the female ovule can only be X. The one carried by pollen of the male plant may be either X or Y. From the pollen, the embryo has a 50/50 chance of receiving an X, likewise for Y; hance, male and female progeny appear in equal numbers (in humans, the sperm carries either an X or a Y chromosome.)
MALE PLANT FLOWERING
Under natural light, males usually start to flower from one to four weeks before the females. Where the photoperiod is artificially controlled, as with electric lights, males respond quickly (in about a week) to a change to short photoperiods and usually show flowers sooner than the females.
In the week or two prior to flowering and throughout flowering, many common marijuana varieties follow two general growth patterns which depend on gender. With these varieties, you can tell gender by the spacing between the leaves (internodes). For the female, the emphasis is on compact growth. Each new leaf grows closer to the last, until the top of the plant is obscured by tightly knit leaves. The male elongates just prior to showing flowers. New growth is spaced well apart and raises the male to a taller stature. This may by the first time the male shows its classic tall, loosely arranged profile. The male preflower may be described as a “ball on a stick.” However, its most recognizable feature is its absence of pistils. Sometimes, a male plant will develop mature staminate flowers after prolonged periods of vegetative growth. These appear in clusters around the nodes. The following image shows a male plant in early flowering. Staminate flowers are located at the node between the stipule and emerging branch.
Male flowers develop quickly, in about one to two weeks on a vigorous plant, not uniformly. Scattered flowers may open a week or more before and after the general flowering, extending the flowering stage to about four weeks.
The flowering stage continues to demonstrate the male’s tall, relatively sparse growth. Most of the flowers develop near the top of the plant, well above the shorter females. The immature flower buds first appear at the tips of the main stem and branches. Then tiny branches sprout from the leaf axils, bearing smaller clusters of flowers. The immature male flowers are closed, usually green, and develop in tight clusters of knob-like buds. The main parts of the male flowers are five petal-like sepals which enclose the sexual organs. As each flower matures, the sepals open in a radiating pattern to reveal five pendulous anthers (stamens).
Inside the ovoid, sac-shaped anthers, pollen grains develop. Initially, pollen sifts through two pores near the top of the anther; then, starting from the pores, longitudinal slits slowly open (zipperlike) over the course of a day, releasing pollen to the wind. Once a flower sheds pollen, it shortly dies and falls from the plant. Normally, male plants begin to die one to two weeks after the bulk of their flowers have shed pollen. Healthy males may continue to flower for several more weeks, but secondary growth seldom has the vigour of initial bloom.
The female plant generally starts to flower later than the male, under either natural light or an artificially controlled photoperiod. Female marijuana plants flower when the average daily photoperiod is less then about 12 to 13 hours. However, some varieties and individuals may flower with a photoperiod of over 14 hours. Some Colombian varieties may not respond until the photoperiod falls below 12 hours for a period of up to three weeks.
The duration of flowering also depends on the particular rhythm of the variety, as well as growing conditions, and whether or not the plant is pollinated. Within these variables, females maintain vigorous growth and continue to rapidly form flowers for a period that ranges from 10 days to about eight weeks.
Females generally do not grow much taller during flowering. Growth emphasises a “filling out,” as flower clusters develop from each leaf axil and growing tip. Normally, the flowers arise in pairs, but the pairs form tight cluster of 10 to over 100 individual flowers that are interspersed with small leaves. These clusters are the “buds” of commercial marijuana. Along the top of the main stem and vigorous branches, “buds” may form so thickly that the last foot or more of stem is completely covered. Usually the leaves that accompany the flowers tend toward simpler structure, until each leaf has one to three blades.
Bracts are covered more densely with large resin glands than is any other part of the plant, and are the most potent part of the harvest. Resin glands may also be seen on the small leaves that are interspersed among the flowers.
The differences between male and female Cannabis become more apparent as the plants mature. The same can be said of the differences between varieties. Often, two varieties may appear to be similar, until they actually flowers and fill out to different forms. These appear in many ways: some varieties maintain opposite phyllotaxy with long internodes throughout flowering; bud sizes vary from about one-half inch to about three inches, with a norm of about one to two inches; buds may be tightly arranged along the stem, yielding a “cola” two feet long and four inches thick; and some varieties only form buds along their main stem and branch tips, with a few “buds” forming along the branches.
When a female is well-pollinated, growth slows and the plant’s energy goes into forming seeds and thus into the continuation of the species. Some plants (but only the more vigorous ones) will renew flowering even when pollinated. Females that are not well-pollinated continue to form flowers rapidly. This extends the normal flowering period, of 10 days to four weeks, up to eight weeks or more.
Individual flowers are pollinated by individual pollen grains. In a matter of minutes from its landing on a stigma, the pollen grain begins to grow a microscopic tube, which penetrates the stigma and reaches the awaiting ovule wrapped within the bracts. The pollen tube is a passageway for the male’s genetic contributions to the formation of the embryo (seed).
The union of the male and female complements of genes completes fertilisation and initiates seed formation. The stigmas, having served their purpose, shrivel and die, turning rust or brown colour. On a vigorous female, the seeds reach maturity in about 10 days. When growing conditions are poor, the seed may take five weeks to ripen to full size and colour. Naturally, all the flowers do not form, nor are they pollinated at the same time – and there will be seeds that reach maturity weeks before others do. Although each flower must be individually fertilised to produce a seed, a single male plant can release many millions of pollen grains. A large female plant can produce over 10,000 seeds.
Sexual Variants in Cannabis
Cannabis has been studied for many years because of its unusual sexuality. Besides the normal dioecious pattern, where each plant bears exclusively male or female flowers, it is not uncommon for some plants to have both male and female flowers. These are called hermaphrodites, or monoecious plants, or intersexes. Hermaphroditic plants form normal flowers of both sexes in a wide variety of arrangements, in both random and uniform distributions.
Some hermaphrodites seem to be genetically determined (protogenous). That is, they naturally form flowers of both sexes given normal growing conditions. Possibly genes carried on the autosomes (the chromosomes other than the sex chromosomes) modify the normal sexual expression. Monoecious varieties have been developed by hemp breeders in order to ensure uniform harvests.
It is also possible that these particular are polyploid, which means they have more than the usual two sets of chromosomes. This kind of hermaphrodite may have XXY (triploid), or XXYY or XXXY (tetraploid) sex chromosomes. However, no naturally occurring polyploids have ever been verified (by observation of the chromosomes) in any population of Cannabis. Polyploids have been induced in Cannabis by using mutagens, such as the alkaloid colchicine.
Whatever then genetic explanation may be, one or more of these natural hermaphrodites may randomly appear in any garden. They are sometimes faster-maturing, have larger leaves, and are larger in overall size than their unisexual siblings. They usually form flowers of both sexes uniformly in time and distribution, and in some unusual patterns. For example, from Mexican seed, we have seen a plant on which separate flowering cluster consisted of both female and male flowers: and upper section of female flowers had upraised stigmas, and a lower section of male flowers dangled beneath the female flowers. In other plants from Mexican seed, the growing tips throughout the plant have female flowers; male flowers sprout from the leaf axils along the main stem and branches. Plants from “Thai” seed sometimes form male and female flowers on separate branches. Branches with female flowers tend to predominate, but branches having mostly male flowers are located throughout the plant.
Abnormal Flowers, Intersexes, Reversals
Gender is set in the new plant at the time of fertilisation by its inheritance of either the X or the Y chromosome from the male (staminate) plant. With germination of the seed, the environment comes into play. Heritage sets the genetic program, but the environment can influence how the program runs. (Sexual expression in Cannabis is delicately balanced between the two.) The photoperiod, for example, controls the plant’s sequence of development. Also, the plant’s metabolism and life processes are dependent on growing conditions. When the environment does not allow a balance to be maintained, the normal genetic program may not be followed. This is mirrored by abnormal growth or sexual expression.
Abnormal Flower Abnormal sexual expression includes a whole range of possibilities. Individual flowers may form abnormally, and may contain varying degrees of both male and female flower parts. For instance, a male flower may bear a stigma; or an anther may protrude from the bracts of a female flower. Abnormally formed flowers are not often seen on healthy plants, although if one looks hard enough, a few may be found in most crops. When many of the flowers are abnormal, an improper photoperiod (coupled with poor health) is the most likely cause. Abnormal flowers sometimes form on marijuana grown out of season, such as with winter or spring crops grown under natural light.
Intersexes and Reversals Much more common than abnormally formed flowers is for the plant’s sex to be confused. One may find an isolated male flower or two; or there may be many clusters of male flowers on an otherwise female plant, or vice versa. These plants are called intersexes (also hermaphrodites or monoecious plants). Intersexes due to environment causes differ from natural hermaphrodite in having random distributions and proportions of male and female flowers. In more extreme cases, a plant may completely reverse sex. For example, a female may flowers normally for several weeks, then put forth new, sparse growth, typical of the male, on which male flowers develop. The complete reversal from male flowering to female flowering also happens.
All other things being equal, the potency of intersexes and reversed plants is usually less than that of normal plants. If there are reversals or intersexes, both of the sexes will usually be affected. Female plants that reverse to male flowering show the biggest decline. Not only is the grass less potent, but the amount of marijuana harvested from male flowers is negligible compared to the amount of marijuana that can be harvested from a normal female. Plants that change from male to female flowering usually increase their potency, because of the growth of female flower bracts with their higher concentration of resin. Female flowers on male plants seldom form as thickly or vigorously as on a normal female. Between the loss in potency and the loss in yield because of females changing to males, a crop from such plants is usually inferior, in both yield and potency, to one from normal plants.
Many environmental factors can cause intersexes and sexual reversals. These include photoperiod, low light intensity, applications of ultraviolet light, low temperatures, mutilation or severe pruning, nutrient imbalances or deficiencies, senescence (old age), and applications of various chemicals (see bibliography on sex determination).
The photoperiod (or time of planting using natural light) is the most important factor to consider for normal flowering. In 1931, J. Schaffner (105) showed that the percentage of hemp plants that had confused sexual characteristics depended on the time of year they were planted. Normal flowering (less than five percent of the plants are intersexes) occurred when the seeds were sown in May, June, or July, the months when the photoperiod is longest and light intensity is strongest. When planted sooner or later in the year, the percentage of intersexuals increased steadily, until about 90 percent of the plants were intersexual when planted during November or early December.
Marijuana plants need more time to develop than hemp plants at latitudes in the United States. Considering potency, size, and normal flowering, the best time to sow for the summer crop is during the month of April. Farmers in the south could start the plants as late as June and still expect fully developed plants.
If artificial light is used, the length of the photoperiod can influence sexual expression. Normal flowering, with about equal numbers of male and female plants, seems to occur when the photoperiod is from 15 to 17 hours of light for a period of three to five months. The photoperiod is then shortened to 12 hours to induce flowering. With longer photoperiods, from 18 to 24 hours a day, the ratio of males to females changes, depending on whether flowering is induced earlier or later in the plant’s life. When the plants are grown with long photoperiods for six months or more, usually there are at least 10 percent more male then female plants. When flowering is induced within three months of age, more females develop. Actually, the “extra” males or females are reversed plants, but the reversals occur before the plants flower in their natural genders.
Some plants will flower normally without a cutting of the photoperiod. But more often, females will not form thick buds unless the light cycle is cut to a period of 12 hours duration. Don’t make the light cycle any shorter than 12 hours, unless the females have not shown flowers after three weeks of 12-hour days. Then cut the light cycle to 11 hours. Flowers should appear in about one week.
Anytime the light cycle is cut to less than 11 hours, some intersexes or reversed plant usually develop. This fact leads to a procedure for increasing the numbers of female flowers indoors. The crops can be grown for three months under a long photoperiod (18 or more hours of light). The light cycle is then cut to 10 hours. Although the harvest is young (about five months) there will be many more female flower buds than with normal flowering. More plants will develop female flowers initially, and male plants usually reverse to females after a few weeks of flowering.
Of the other environmental factors that can affect sexual expression in Cannabis, none are as predictable as the photoperiod. Factors such as nutrients or pruning affect the plant’s overall health and metabolism, and can be dealt with by two general thoughts. First, good growing conditions lead to healthy plants and normal flowering: female and male plants occur in about equal numbers, with few (if any) intersexes or reversed plants. Poor growing conditions lead to reduced health and vigour, and oftentimes to confused sex in the adult plant. Second, the age of the plants seems to influence reversals. Male plants often show female flowers when the plant is young (vigorous) during flowering. Females seven or more months old (weaker) often develop male flowers after flowering normally for a few weeks.
Anytime the plant’s normal growth pattern is disrupted, normal flowering may be affected. For instance, plant propagated from cuttings sometimes reverse sex, as do those grown for more than one season.
Sexing the Plants
The female plant is more desirable than the male for marijuana cultivation. The female flowering clusters (bus) are usually the most potent parts of the harvest. Also, given room to develop, a female generally will yield twice as much marijuana as her male counterpart. More of her weight consists of top-quality buds.
Because the female yields marijuana in greater quantity and sooner you can devote your attention to nurturing the females. Where space is limited, such as in indoor gardens and small outdoor plots most growers prefer to remove the males as soon as possible, and leave all available space for the females. To harvest sinsemilla (seedless female buds), you must remove the male plants before they mature and release pollen.
Differences in the appearance of male and female Cannabis become more apparent toward maturation. During the seedling stage, gender is virtually impossible to distinguish, although in some varieties the male seedling may appear slightly taller and may develop more quickly.
We know of no way to discover gender with any certainty until each plant actually forms either pollen-bearing male flowers or seed-bearing female flowers. However, certain general characteristics may help. Using guidelines like the following, growers who are familiar with a particular variety can often predict gender fairly accurately by the middle stage of the plant’s life.
Early Vegetative Growth
After the initial seedling stage, female plants generally develop more complex branching than the male. The male is usually slightly taller and less branched. (Under artificial light, the differences in height and branching are less apparent throughout growth.)
Some plants develop a marked swelling at the nodes, which is more common and pronounced on female plants.
Middle Vegetative Growth
In the second to fourth months of growth, plants commonly form a few isolated flowers long before the actual flowering stage begins. These premature flowers are most often found between the eighth and twelfth nodes on the main stem. Often they appear near each stipule (leaf spur) on several successive nodes, at a distance two to six nodes below the growing tip. These individual flowers may not develop fully and are often hard to distinguish as male or female flowers. The fuzzy white stigmas of the female flower may not appear, and the male flowers seldom opens but remains a tightly closed knob. However, the male flower differs from the female; it is raised on a tiny stalk, and the knob is symmetrical. The female flower appear stalkless and more leaflike.
The presence of premature female flowers does not assure that the plant is a female, but premature male flowers almost always indicate a male plant. Unfortunately, it is much less common for male plants to develop premature male flowers than for female flowers to appear on either plant. For example, in one garden of 25 mixed-variety plants, by age 14 weeks, 15 plants showed well-formed, premature female flowers with raised stigmas. Eight of these plants matured into females and seven became males. Only two plants showed premature male flowers and both of these developed into males. The eight remaining plants did not develop premature flowers or otherwise distinguishable organs until the actual flowering stage at the age of 21 weeks. From these eight, there were four females, three males, and one plant bearing both male and female flowers (hermaphrodite). It does seem, however, that plants bearing well-formed female flowers, on several successive node, usually turn out to be females.
PART 2: Life Cycles of Cannabis……………………..
An Advanced Study: The Propagation and Breeding of Distinctive Cannabis
by Robert Connell Clarke
Sinsemilla Life Cycle of Cannabis
Cannabis is a tall, erect, annual herb. Provided with an open sunny environment, light well-drained composted soil, and ample irrigation, Cannabis can grow to a height of 6 meters (about 20 feet) in a 4-6 month growing season. Exposed river banks, meadows, and agricultural lands are ideal habitats for Cannabis since all offer good sunlight. In this example an imported seed from Thailand is grown without pruning and becomes a large female plant. A cross with a cutting from a male plant of Mexican origin results in hybrid seed which is stored for later planting. This example is representative of the outdoor growth of Cannabis in temperate climates.
Seeds are planted in the spring and usually germinate in 3 to 7 days. The seedling emerges from the ground by the straightening of the hypocotyl (embryonic stem). The cotyledons (seed leaves) are slightly unequal in size, narrowed to the base and rounded or blunt to the tip. The hypocotyl ranges from 1 to 10 centimeters (1A to 3 inches) in length. About 10 centimeters or less above the cotyledons, the first true leaves arise, a pair of oppositely oriented single leaflets each with a distinct petiole (leaf stem) rotated one-quarter turn from the cotyledons. Subsequent pairs of leaves arise in opposite formation and a variously shaped leaf sequence develops with the second pair of leaves having 3 leaflets, the third 5 and so on up to 11 leaflets. Occasionally the first pair of leaves will have 3 leaflets each rather than 1 and the second pair, 5 leaflets each.
If a plant is not crowded, limbs will grow from small buds (located at the intersection of petioles) along the main stem. Each sinsemilla (seedless drug Cannabis) plant is provided with plenty of room to grow long axial limbs and extensive fine roots to increase floral production. Under favorable conditions Cannabis grows up to 7 centimeters (21A inches) a day in height during the long days of summer.
Cannabis shows a dual response to daylength; during the first two to three months of growth it responds to increasing daylength with more vigorous growth, but in the same season the plant requires shorter days to flower and complete its life cycle.
LIFE CYCLE OF CANNABIS I Juvenile Stage
Cannabis flowers when exposed to a critical daylength which varies with the strain. Critical daylength applies only to plants which fail to flower under continuous illumination, since those which flower under continuous illumination have no critical daylength. Most strains have an absolute requirement of inductive photoperiods (short days or long nights) to induce fertile flowering and less than this will result in the formation of undifferentiated primordia (unformed flowers) only.
The time taken to form primordia varies with the length of the inductive photoperiod. Given 10 hours per day of light a strain may only take 10 days to flower, whereas if given 16 hours per day it may take up to 90 days. Inductive photoperiods of less than 8 hours per day do not seem to accelerate primordia formation. Dark (night) cycles must be uninterrupted to induce flowering (see appendix).
Cannabis is a dioecious plant, which means that the male and female flowers develop on separate plants, although monoecious examples with both sexes on one plant are found. The development of branches containing flowering organs varies greatly between males and females: the male flowers hang in long, loose, multi-branched, clustered limbs up to 30 centimeters (12 inches) long, while the female flowers are tightly crowded between small leaves.
Note: Female Cannabis flowers and plants will be referred to as pistillate and male flowers and plants will be referred to as staminate in the remainder of this text. This convention is more accurate and makes examples of complex aberrant sexuality easier to understand.
The first sign of flowering in Cannabis is the appearance of undifferentiated flower primordia along the main stem at the nodes (intersections) of the petiole, behind the stipule (leaf spur). In the prefloral phase, the sexes of Cannabis are indistinguishable except for general trends in shape.
When the primordia first appear they are undifferentiated sexually, but soon the males can be identified by their curved claw shape, soon followed by the differentiation of round pointed flower buds having five radial segments. The females are recognized by the enlargement of a symmetrical tubular calyx (floral sheath). They are easier to recognize at a young age than male primordia. The first female calyxes tend to lack paired pistils (pollen-catching appendages) though initial male flowers often mature and shed viable pollen. In some individuals, especially hybrids, small non-flowering limbs will form at the nodes and are often confused with male primordia.
Cultivators wait until actual flowers form to positively determine the sex of Cannabis
The female plants tend to be shorter and have more branches than the male. Female plants are leafy to the top with many leaves surrounding the flowers, while male plants have fewer leaves near the top with few if any leaves along the extended flowering limbs.
*The term pistil has developed a special meaning with respect to Cannabis which differs slightly from the precise botanical definition. This has come about mainly from the large number of cultivators who have casual knowledge of plant anatomy but an intense interest in the reproduction of Cannabis. The precise definition of pistil refers to the combination of ovary, style and stigma. In the more informal usage, pistil refers to the fused style and stigma. The informal sense is used throughout the book since it has become common practice among Cannabis cultivators.
The female flowers appear as two long white, yellow, or pink pistils protruding from the fold of a very thin membranous calyx. The calyx is covered with resin exuding glandular trichomes (hairs). Pistillate flowers are borne in pairs at the nodes one on each side of the petiole behind the stipule of bracts (reduced leaves) which conceal the flowers. The calyx measures 2 to 6 millimeters in length and is closely applied to, and completely contains, the ovary.
In male flowers, five petals (approximately 5 millimeters, or 3/16 inch, long) make up the calyx and may be yellow, white, or green in color. They hang down, and five stamens (approximately 5 millimeters long) emerge, consisting of slender anthers (pollen sacs), splitting upwards from the tip and suspended on thin filaments. The exterior surface of the staminate calyx is covered with non-glandular trichomes. The pollen grains are nearly spherical slightly yellow, and 25 to 30 microns (p) in diameter. The surface is smooth and exhibits 2 to 4 germ pores.
Before the start of flowering, the phyllotaxy (leaf arrangement) reverses and the number of leaflets per leaf decreases until a small single leaflet appears below each pair of calyxes. The phyllotaxy also changes from decussate (opposite) to alternate (staggered) and usually remains alternate throughout the floral stages regardless of sexual type.
The differences in flowering patterns of male and female plants are expressed in many ways. Soon after dehiscence (pollen shedding) the staminate plant dies, while the pistillate plant may mature up to five months after viable flowers are formed if little or no fertilization occurs. Compared with pistillate plants, staminate plants show a more rapid increase in height and a more rapid decrease in leaf size to the bracts which accompany the flowers. Staminate plants tend to flower up to one month earlier than pistillate plants; however, pistillate plants often differentiate primordia one to two weeks before staminate plants.
Many factors contribute to determining the sexuality of a flowering Cannabis plant. Under average conditions with a normal inductive photoperiod, Cannabis will bloom and produce approximately equal numbers of pure staminate and pure pistillate plants with a few hermaphrodites (both sexes on the same plant). Under conditions of extreme stress, such as nutrient excess or deficiency, mutilation, and altered light cycles, populations have been shown to depart greatly from the expected one-to-one staminate to pistillate ratio.
Just prior to dehiscence, the pollen nucleus divides to produce a small reproductive cell accompanied by a large vegetative cell, both of which are contained within the mature pollen grain. Germination occurs 15 to 20 minutes after contact with a pistil. As the pollen tube grows the vegetative cell remains in the pollen grain while the generative cell enters the pollen tube and migrates toward the ovule. The generative cell divides into two gametes (sex cells) as it travels the length of the pollen tube.
Pollination of the pistillate flower results in the loss of the paired pistils and a swelling of the tubular calyx where the ovule is enlarging. The staminate plants die after shedding pollen. After approximately 14 to 35 days the seed is matured and drops from the plant, leaving the dry calyx attached to the stem. This completes the normally 4 to 6 month life cycle, which may take as little as 2 months or as long as 10 months. Fresh seeds approach 100% viability, but this decreases with age.
The hard mature seed is partially surrounded by the calyx and is variously patterned in grey, brown, or black. Elongated and slightly compressed, it measures 2 to 6 millimeters (1/16 to 3/16 inch) in length and 2 to 4 millimeters (1/16 to 1/8 inch) in maximum diameter.
Careful closed pollinations of a fewselected limbs yield hundreds of seeds of known parentage, which are removed after they are mature and beginning to fall from the calyxes. The remaining floral clusters are sinsemilla or seedless and continue to mature on the plant. As the unfertilized calyxes swell, the glandular trichomes on the surface grow and secrete aromatic THC-laden resins. The mature, pungent, sticky floral clusters are harvested, dried, and sampled. The preceding simplified life cycle of sinsemilla Cannabis exemplifies the production of valuable seeds without compromising the production of seedless floral clusters.