<\/div>\n\n\n\n
Unforgettable Qualities of Cannabis<\/h3>\n\n\n\n Cannabis has a unique set of qualities that influence its life cycle and reproduction. Working with these natural qualities is essential to breeding cannabis. Please remember them when working on all breeding projects.<\/p>\n\n\n\n
Cannabis is photoperiodic reactive<\/strong>, flowering under 12 hours of uninterrupted darkness and 12 hours of light, which gives breeders the ability to grow up to 6 crops every year. These characteristics facilitate fast-track breeding because 6 years of crosses can be completed in 1 year.<\/p>\n\n\n\nCannabis is dioecious<\/strong>; it produces each of the male (staminate; stamen-bearing) and female (pistillate; pistil-bearing) sexual organs on different individuals, and it is one of the few annual dioecious plants. This quality makes it very easy to cross an individual male cannabis plant with an individual or population of females.<\/p>\n\n\n\nMale (staminate) and female (pistillate) plants are easy to distinguish. See close-up details of male and female flowers along with full descriptions below in this chapter.<\/p>\n\n\n\n
Monoecious<\/strong> varieties produce both staminate and pistillate flowers on the same plant. Monoecious varieties are mainly used for hemp seed production. Monoecious varieties do not make good medical cannabis breeding stock. Plants with both male and female flowers are often called by the misnomer “hermaphrodite.”<\/p>\n\n\n\nOutcrossing plant: <\/strong>Outcrossing cannabis grows best when plants cross with other plants with different genes. Corn, dogs, cannabis, and people are all outcrossers. Backcrossing and inbreeding cannabis for too many generations will be detrimental.<\/p>\n\n\n\nIntersexuality<\/strong> occurs when flowers of a male plant grow on a predominantly female plant, or when female flowers grow on a predominantly male plant. It is a trait that can be caused by both genetic and environmental factors. Intersex plants with the inherited gene grow flowers from both sexes on the same plant even in perfect growing conditions. Intersex plants are often called by the misnomer “hermaphrodite.”<\/p>\n\n\n\nIndoors, plants are easily exposed to stress\u2014inconsistent or extreme temperature, light cycles, fertilizer, pH, etc.\u2014and this stress can cause female plants to grow an occasional male flower. Breeders prefer not to perpetuate the intersex gene, and when possible, they eliminate it. A single male flower on a female plant can pollinate a big part of the female. See “Feminized Seeds,” on page 521.<\/p>\n\n\n\n
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Physiology of Male and Female Flowers<\/h2>\n\n\n\nMale Flowers<\/h3>\n\n\n\n Male (staminate) cannabis flowers are about 0.25 to 0.5 inches (0.6\u20131.3 cm) long, and thousands of individual flowers can develop on a large plant. Most of the flowers develop in loose clusters (cymes or cymose panicles) of about 5 or 10 flowers each, borne on tiny branches and their side (lateral) branches. The clusters can pile atop each other to form dense aggregates of hundreds of individual flowers, particularly at the ends of stems and branches.<\/p>\n\n\n\n
Each male flower’s calyx consists of 5 tepals (sometimes identified as “sepals”)\u2014usually white, yellowish, or greenish, but often tinged purple\u2014that might be described as “petals,” and 5 pendulous stamens that bear pollen in sacks called anthers. Anthers hang by a thin, threadlike filament, and together, filament and anther make up the stamen. Once mature, 2 openings on opposite sides of each anther open zipper-like, starting at their base, to slowly release their pollen into the wind, carrying it (hopefully) to stigmas. It has been estimated that the thousands of flowers on a single male can release more than 500 million pollen grains.<\/p>\n\n\n\n
Unopened male flower clusters remind some growers of tiny grape clusters, and fresh anthers look somewhat like bunches of tiny bananas. Male flowers are simply called male flowers or male flower clusters, and the pollen holders are referred to as either stamens or anthers.<\/p>\n\n\n\n
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<\/figure><\/div>\n\n\n<\/div>\n\n\n\n
This drawing shows the main parts of a male cannabis plant.<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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‘Jack Herer’ male (staminate) flowers fully formed in clusters but not yet open. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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Outer tepals on the ‘Skunk #1’ staminate calyxes have separated, exposing the pollen-containing anthers. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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As anthers on this ‘Jack Herer’ mature, they rupture to disperse superfine grains of pollen into the air. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n\n
Female Flowers<\/h3>\n\n\n\n Each female marijuana flower has 2 stigmas that protrude from the single ovule enclosed in the bracts; fresh stigmas are “fuzzy” (hirsute), about 0.25 to 0.5 inches (0.6\u20131.3 cm) long, and usually white, but sometimes they are yellowish or pink to red, and, rarely, lavender to purple. Stigmas (stigmata is another botanical plural) are the pollen catchers. Stigmas are often misidentified as pistils. By definition, a pistil is all of the reproductive parts of a flower\u20142 stigmas and an ovule make up the female pistil. Each flower then has only 1 pistil but has 2 stigmas. The term is misused in much of popular culture, which describes a single cannabis flower as having 2 pistils.<\/p>\n\n\n\n
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<\/figure><\/div>\n\n\n<\/div>\n\n\n\n
Staminate anthers on this ‘Skunk #1’ grown in 1987 continue to split open and disperse more and more pollen into the air. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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Anthers hang in the wind after dispersing all their pollen throughout several days. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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Pollen from this ‘Skunk #1’ has been dispersed from anthers in the foreground. Staminate calyxes in the background will disperse pollen in the near future. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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Compare this tiny male flower to the size of a US penny. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n\n
Stigmas begin dying after pollination and begin to turn rust-colored about 3 days later; if not pollinated, as with sinsemilla (seedless buds), stigmas begin to die when they are about 4 or 5 weeks old. Upon landing on a stigma, a pollen grain germinates and begins to grow a pollen tube through the style passageway to pass its DNA to join the ovule\u2019s DNA (the 2 stigmas of each female flower make up a bifid style). The fertilized ovule becomes a fruit, essentially a single seed (an achene). The perianth, which includes the calyx, tightly clasps the seed and often contains tannins, which give mature seeds their mottled or spotted coat. Between a thumb and finger you can rub the perianth off of seeds. A well-pollinated single bud develops dozens of seeds, a cola easily holds many hundreds, and even a small, but thoroughly pollinated female can bear thousands of seeds.<\/p>\n\n\n\n
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This drawing shows the main parts of a female cannabis plant.<\/em><\/p>\n\n\n\n<\/div>\n\n\n\n
Each female flower has a single ovule partially enclosed in its perianth, which is encapsulated by bracteoles, which are covered by a whorl of bracts. The bracts and bracteoles are small, modified leaves that enclose and protect the seed in what some growers refer to as the seedpod.<\/p>\n\n\n\n
Bracts contain the highest concentration of THC and other cannabinoids of any plant part and about 50 percent of a plant\u2019s total THC. The perianth and its calyx contain no THC.<\/p>\n\n\n\n
By definition, a perianth consists of a corolla and a calyx. In more familiar showy flowers, the corolla is the brightly colored petals we generally appreciate when looking at flowers, and the calyx is the smaller green cup (sepals) at the flower\u2019s base. Bright showy colors, large flower sizes, and enticing fragrances have naturally evolved to attract insects such as bees and flies, or animals such as birds and bats that collect and transfer pollen (unintentionally) to other flowers. Cannabis flowers are not brightly colored, large, or enticingly fragrant (at least to most nonhumans); cannabis plants are wind-pollinated with no need to attract insects or animals to carry males\u2019 pollen to female flowers; cannabis plant parts never naturally evolved into colorful, attractive, or showy parts. Cannabis breeders, though, do breed for fragrance and color once cannabinoid content is firmly established.<\/p>\n\n\n\n
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The seed bract still covers the perianth, pistillate calyx, gametes, and ovule connected to a pair of stigmas. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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The seed bract has been stripped away to reveal the perianth and pistillate calyx, both of which appear transparent, covering the gametes and ovule. Note the white stigmas have not been pollinated. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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Stigmas start dying back as soon as pollen is ushered down the shaft to unite with the ovule below. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n\n
The cannabis perianth is only about 6 cells thick, so to distinguish calyx cells from corolla cells is best left to botanists. This book uses the botanically correct term bracts for the green or purple, resin-gland-studded specialized “leaves” encasing each female flower, and uses perianth or calyx for the translucent “veil” that clasps and covers about 60 to 90 percent of a mature seed.<\/p>\n\n\n\n
Hopefully, when growers use botanical terms such as calyx, bracts, stigma, pistil, anther and stamen, they will follow this book and use the terms correctly. Since readers will find seed catalogs and Internet sites calling bracts calyxes and stigmas pistils, it is important for readers to understand this confusion when reading other sources. Hopefully, this chapter will help get us all on the same page.<\/p>\n\n\n\n
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The stigmas on this ‘Haze’ \u00d7 ‘Northern Lights’ \u00d7 ‘Sensi Star’ flower top are just starting to turn a rusty color. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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The perianth on this ‘Skunk #1’ can be seen near the top of the seed on the left. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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The female perianth is clearly visible as a nearly transparent layer covering the seed. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n\n
Sexual Propagation<\/h2>\n\n\n\n Sexual propagation is the process in which male and female sex cells (gametes) from separate parents unite in the female plant to form what will eventually mature into a new, genetically distinct individual. This process occurs when pollen from a male (staminate) parent unites with an ovule within the ovary of a female flower to create an embryo. This embryo, when mature and fully developed, will become a seed.<\/p>\n\n\n\n
In nature, cannabis is wind-pollinated. Male flowers shed pollen (dehiscence), dispersing millions of grains into the wind. Wind carries the pollen to a “chance” rendezvous and acceptance by a female stigma.<\/p>\n\n\n\n
Pollination occurs when male pollen grains land on a female stigma. The evolutionary attraction is both physical and chemical. The grain of pollen, with moisture found in the stigma, germinates. This is the best part: A grain of pollen germinates just like a seed, sending a taproot down, but instead of sending it into the ground, the grain of pollen sends the “root” down the stigma toward the ovary. Once united with the ovary, the pollen fertilizes the ovule. This union creates an embryo that grows within a seed coat. When mature in 4 to 6 weeks, the seed can be planted.<\/p>\n\n\n\n
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Fertilization occurs when the tiny grain of male pollen sticks to the stigma. Then it develops a tube through the style and releases 2 gametes, 1 to fertilize the ovule and 1 to fertilize the endosperm (double fertilization). Seeds are the result of this sexual propagation and contain genetic characteristics of both parents. Once fertilized with male pollen, female plants put the bulk of their energy into producing strong, viable seeds.<\/p>\n\n\n\n
Actual fertilization takes place when the minute grain of male pollen sticks to the stigma. The successful angiosperm pollen grain (gametophyte) containing the male gametes (sperm) gets transported to the stigma, where it germinates and its pollen tube grows down the style to the ovary. Its 2 gametes travel down the tube to where the gametophyte(s) containing the female gametes are held within the carpel. One nucleus fuses with the polar bodies to produce the endosperm tissues, and the other with the ovum to produce the embryo hence the term double fertilization.<\/p>\n\n\n\n
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Close-up photo of a female stigma shows that no resin glands are located on the entire length. The well-defined protruding growth appears as fuzz on the stalk of the stigma. The stigma is the plant version of a vagina. It is covered with stigmatic fluid that acts like glue when a piece of pollen lands on it. The fluid is packed with sugars that are food for the pollen. When in place, the grains of pollen start growing a new \u201cpollen tube,\u201d a long tunnel that pushes through the tissue of the style all the way to the ova, where it fuses with egg cells to create a little baby plant. The ovules go through a series of steps known as meiosis, a type of cell division by which a cell duplicates into 2 genetically identical daughter cells. The chromosomes in the cell nucleus separate into 2 matching sets of chromosomes, each with a nucleus.<\/p>\n\n\n\n
Deoxyribonucleic acid (DNA) or “genetic material”* is coiled into long strands or chromosomes. The DNA is located inside the nucleus of each cell. When cannabis is pollinated, each individual seed inherits 10 different chromosomes from the male, and 10 different chromosomes from the seed mother\u201420 chromosomes total. Each seed has 2 copies of each of the 10 chromosomes, or 1 full genome each. There are 2 copies of every gene in the plant, 1 from the mother and 1 from the father. Every cell in the plant has a copy of this unique DNA. The genetic code of this unique individual is embedded in a specific location along the length of the chromosome strands.<\/p>\n\n\n
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Each seed<\/strong> contains genes from both parents. Progeny grown from seed usually have slightly different traits than other plants from the same seed lot. The same happens in humans; biological children are different from one another in many aspects and at the same time they resemble their parents. In cannabis, the variability is marked, like it is in apples.<\/p>\n\n\n\nSexual reproduction is used to cross different individuals with a population or family of plants. It can also be used to hybridize unrelated lines and inbreed their offspring. This phenomenon, \u201crecombination of traits,\u201d also gives breeders the opportunity to recover individuals with a combination of the positive traits of both parental lines.<\/p>\n\n\n\n
Genes are hereditary units that consist of a sequence of DNA that resides in a precise place on a chromosome and it determines a specific trait in cannabis. Little bits of DNA are codes or templates for proteins.<\/p>\n\n\n\n
Proteins are made in the sequence of DNA. Like instructions in a recipe for making brownies, the DNA and sequence of proteins is the recipe or instructions.<\/p>\n\n\n\n
Having 2<\/strong> versions of the same protein<\/strong>, from 2 different genes, is better than having just 1, particularly if the protein plays some vital part in cannabinoid production. This effect is called overdominance. For example, if there are 2 different proteins, and both work well but 1 works a little better under hot conditions and the other works well under cool conditions. Having 2 versions of the same protein gives the plant a wider range of climates where it produces effectively. See \u201cMultiline,\u201d on page 531.<\/p>\n\n\n\nThe majority of DNA is the same; it deals with basic cell processes, photosynthesis, chlorophyll production, etc. A few or a combination of genes control variables such as height, leaf shape, fragrance, and disease resistance. But we are not sure exactly which genes are responsible for specific traits, even though the cannabis genome has been mapped. These traits are influenced by multigene families (a group of genes that evolved to become a little bit different from each other, even though they started out as copies of the same gene). Knowing the named genes<\/strong> would make it easier to find individual plants with your desired traits. But single genes controlling specific traits of cannabis are neither isolated nor well studied.<\/p>\n\n\n\nMultigene traits allow you to fine-tune to your favorite characteristics. For example, a single gene that controls leaf size would give only 2 leaf sizes, large and small. Many genes influencing the same trait provide many different leaf sizes.<\/p>\n\n\n\n
Naturally occurring mutated cannabis genes<\/strong> are uncommon. They are abnormal genes that are mutations of normal genes. When a mutated gene combines with a normal gene, there is no detrimental outcome. But when 2 mutated genes join, the result is much different.<\/p>\n\n\n\nFor example, in people and animals the number of albinos or dwarfs is minimal. The same is true in cannabis. Growing large populations of cannabis or treating cannabis with stress or chemicals will bring about mutation. Overall, most cannabis plants grow normally with no mutations whatsoever. Many different genes control the desirable traits we care about. Broken recessive genes do not play a role in most breeding programs.<\/p>\n\n\n\n
Deleterious recessive genes:<\/strong> The plants most likely to have the same dangerous mutation in the immediate family are inbred. Marry your sister and inbreeding genes take over to start all kinds of problems because recessive deleterious genes appear.<\/p>\n\n\n\n<\/div>\n\n\n
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‘Royal Moby’, predominantly sativa, is very THC-potent. This variety is quite similar to ‘Moby Dick’ from Dinafem Seeds in Spain. Once a good variety reaches the marketplace, many similar varieties appear within a year or two.<\/em><\/p>\n\n\n\n<\/div>\n\n\n\n
Classical Cannabis Breeding<\/h2>\n\n\n\n Classical breeding is an ancient cyclical process that cannabis breeders still use today. Decisions are made based on observation of large numbers of plants; the breeder does not know exactly what genes have been introduced to the new cultivars. All the breeder can do is choose plants based upon visual inspection, smell, and gut-feeling.<\/p>\n\n\n\n
Classical cannabis breeding is simple: 2 varieties, a male and a female, are chosen. Each parent has desirable characteristics\u2014fragrance, potency, mold resistance, and so forth. The male pollen fertilizes the female flower and their genes combine into a new genetic mix contained in the seed.<\/p>\n\n\n\n
The next step is to choose individual plants with the desirable traits of both parents. A lot of the time you get lucky and the offspring carry the desirable genes and traits. Cannabis breeders often take clones of these desirable individual plants. Too often they do not retain the male plant and have a \u201cclone only\u201d variety.<\/p>\n\n\n\n
When a desirable trait has been bred into a plant, crossing other plants to this parent makes new plants similar to the favored parent. For example, to make the mildew-resistant progeny of the cross most like the high-yielding parent, the progeny will be backcrossed to that parent for several generations (see \u201cBackcrossing,\u201d page 519). This process removes most of the genetic contribution of the mildew-resistant parent.<\/p>\n\n\n\n
To breed for mold resistance, grow out plants in moldy conditions. Remove plants from garden that contract mold easily. Keep plants that do not get mold or are late to get mold. Breed plants that do not mold.<\/p>\n\n\n\n
It is very difficult to isolate specific genes to guarantee specific qualities such as extreme resistance to powdery mildew or insect and mite attacks. There are recessive genes and dominant genes that are controlled by alleles; this is where breeding becomes much more complex. See \u201cInfluence of Alleles.\u201d<\/p>\n\n\n\n
Other traits, such as acclimatization to a specific climate, are relatively easy because the plants that grow best in the environment are continually selected. Organic gardeners breed plants acclimated to their outdoor climate and achieve much higher yields. For this reason, organic medical gardeners in Northern California are able to grow 10-pound (4.5 kg) plants.<\/p>\n\n\n\n
To breed for cold tolerance, grow out plants in cold conditions. Remove plants that suffer cold damage easily. Breed plants that withstand cold temperatures.<\/p>\n\n\n\n
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Thin-Layer Chromatography<\/h3>\n\n\n\n Thin-layer chromatography can be used to take tests and make selections based upon cannabinoid profiles. Cannabinoid profiles are similar throughout a plant\u2019s life stages, and breeding decisions can be based upon these profiles. For example, the cannabinoid profile can be tested on 2-month-old seedlings. Plants with desirable profiles are kept, and those with undesirable plants are culled.<\/p>\n\n\n\n
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See \u201cModern Cannabis Breeding\u201d at the end of this chapter for an overview of marker-assisted selection (MAS).<\/p>\n\n\n\n
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Influence of Alleles<\/h2>\n\n\n\n The phenotypes seen in a given individual are the result of an interaction between the plant\u2019s genotype and the environment. For example, here are 3 phenotypes: short, medium, and tall. Remember, the genotype describes the genetic condition responsible for the phenotype, and to represent it in discussion we assign it symbols.<\/p>\n\n\n\n
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Phenotypes<\/strong><\/td>Genotypes<\/strong><\/td><\/tr>short<\/td> ss<\/td><\/tr> medium<\/td> ss<\/td><\/tr> tall<\/td> SS<\/td><\/tr><\/tbody><\/table><\/figure>\n\n\n\n<\/div>\n\n\n\n
There are always 2 versions of every gene (allele). For example, if there are 2 lower-case \u201cs\u201d plants with \u201csmall stature\u201d the plant will be shorter. But if the plant has capital \u201cS\u201d and has the \u201ctall\u201d gene, the phenotype is tall. If both genes are inherited, the plant is medium height.<\/p>\n\n\n\n
Homozygous \/ heterozygous: These are terms used in describing the genotypic condition of a plant, with regard to the similarity of the alleles for a given trait. If a plant is homozygous for a given trait, it has 2 copies of the same allele. If a plant is heterozygous, it has 2 different alleles for a given trait.<\/p>\n\n\n\n
The offspring inherits 1 set of alleles from each parent. This inheritance of alleles can be homozygous (both alleles are the same) or heterozygous (each allele is different). Furthermore, recessive alleles do not surface completely for several generations. The influence of alleles makes it impossible to use simple mathematical probability to predict the outcome of offspring.<\/p>\n\n\n\n
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In August 2011, Dr. Kevin McKernan announced that his company had successfully mapped the genomes (shotgun sequence) for Cannabis sativa (‘Chemdawg’ variety) and later C. indica (‘LA Confidential’). Medicinal Genomics then publicized its work on C. sativa via Amazon\u2019s EC2, a cloud-computing service that gives free access to the scientific community. Search “Cannabis genome EC2 cloud” on www.google.com<\/a> for more information.<\/p>\n\n\n\n \n\n\n\n<\/div>\n\n\n
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This indica-dominant \u2018Peyote Purple\u2019 was developed by CannaBioGen.<\/em><\/p>\n\n\n\n<\/div>\n\n\n\n
Dominant and Recessive Traits<\/h2>\n\n\n\n Dominant and recessive traits are dictated by alleles that are inherited from both parents. But, even though the cannabis genome has been decoded, specific gene function has not been deciphered. Consequently, the examples below must be used as guidelines because many traits are driven by a combination of genes.<\/p>\n\n\n\n
Dominant:<\/strong> An intra-allelic interaction such that the presence of an allele of 1 parent masks the presence of an allele from another parent plant, in the expression of a given trait in the progeny. Only the dominant trait is shown in the first generation of offspring. Of the F2 generation, 75 percent will also show the dominant condition.<\/p>\n\n\n\nRecessive:<\/strong> An intra-allelic interaction such that an allele of 1 parent is masked by the presence of an allele from the other parent plant, in the expression of a given trait in the progeny. The recessive trait is not shown in the first generation of progeny (F1) but will reappear if siblings are mated, and the F2 progeny will result in 25 percent of plants showing the recessive condition.<\/p>\n\n\n\n<\/div>\n\n\n
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\u2018Afghani\u2019-dominant crosses are in the foreground and sativa-dominant crosses are in the background. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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The \u2018Afghani\u2019 seedling on the left demonstrates the dominant traits of short, squat growth and broad leaves. The \u2018Kush\u2019 seedling on the right demonstrates the dominant traits of taller growth, and has much different leaf formation.<\/em><\/p>\n\n\n\n<\/div>\n\n\n
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This genetically purple \u2018Mexican\u2019 sativa from 1980 is one of the varieties that added a purple hue to many current varieties. (MF)<\/em><\/p>\n\n\n\n<\/div>\n\n\n\n
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Numbers Can Be Misleading<\/strong><\/p>\n\n\n\nSimplistic math models are often used to explain what happens when male and female cannabis plants are crossed. These models do not take into account dominant and recessive genes, they allow nothing for genotype and phenotype, and they do not consider that specific traits may be controlled by many different genes.<\/p>\n\n\n\n
Use the Punnett square to help predict the outcome.<\/p>\n\n\n\n
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