Uniform plant-wide flower development??

jockogreen

Active Member
When a plant is in full flowering, and there are buds developing at each node as well as the main cola, do all the buds develop at the same rate? Or do some ripen first?

I don't imagine they all are at the same stage of development, much as fruit on a tree don't all ripen at the same time. But the operative word in that sentence is imagine. I don't know.

Does anybody here?
 

Reiss

Well-Known Member
The whole plant will flower, the buds near the bottom will mature quicker than the buds further up and can usually be harvested earlier too. Depends on how you grow the plant also. I LST which means that most of my buds receive equal light and mature to be pretty much the same through-out.
 

MrFishy

Well-Known Member
On my grows, the plants lower buds are usually still young and vibrant (fluros) when the upper two-thirds is ripe for harvest. I just leave em' (baby buds) if I've the room to spare. They get really resinous with all that growth out of the lights way. In my experience, the bigger, upper buds mature much faster than the lower, shaded stuff.
 

jockogreen

Active Member
So Reiss says the buds mature from the bottom up, Mr. Fishy says the the opposite, top buds mature first. Sounds like lighting may be a factor. My babies are growing outdoors, but I have been 12/12ing them by bringing them inside in the evening; my point is they get get natural sunlight. And the nodes in the middle seem to be maturing the fastest. They're not large, but they're getting orange and red. The upper flowers are more creamy green iciness. And the lower nodes' flowers are even less mature.

Just wondering what other people are seeing (aside from the fact I sometimes fuss a bit too much over my plants. i'm sure i'm not the only one here who does that.)

Or can anyone point me to a good biological resource, if I were to want to find out more about the biology of the flowering process?
 

Reiss

Well-Known Member
As you can tell from the contradicting replies, it depends on many factors. In my experience, the smaller buds near the bottom mature quicker than the fat buds near the top.
best advice is to buy a 30x jewellers loupe off ebay and look at the trichs on your buds.
 

jockogreen

Active Member
Yes yes yes. I already have the Radioshack (tm) pocket 420 microscope. And of course, I'm watching my plants closely. Now I know trich development doesn't necessarily parallel visible bud development, and that buds can turn red while the trichs still remain clear, etc. What I'm really seeking is some more technical information.

What are the changes occurring in the plant? What is going on in terms of auxin signaling and other biochemical processes in the plant? What is causing nutrient uptake to change, and how is it changing? Questions such as these.

I'm geekish. So I wanna know the science behind what's happening.
 

jockogreen

Active Member
I think I found what I was looking for. It was in an e-book I found called Marijuana Botany, An Advanced Study: The Propagation and Breeding of Distinctive Cannabis
by Robert Connell Clarke.


Here's the relevant passage. It's long and technical, but very interesting.

===============

Floral Maturation

The individual pistillate calyxes and the composite floral clusters change as they mature. External changes indicate that internal biochemical metabolic changes are also occurring. When the external changes can be connected with the invisible internal metabolic changes, then the cultivator is in a better position to decide when to har vest floral clusters. With years of experience this becomes intuition, but there are general correlations which can put the process in more objective terms. The calyxes first appear as single, thin, tubular, green sheaths surrounding an ovule at the basal attached end with a pair of thin white, yellowish green, or purple pistils attached to the ovule and protruding from the tip fold of the calyx. As the flower begins to age and mature, the pistils grow longer and the calyx enlarges slightly to its full length. Next, the calyx begins to swell as resin secretion increases, and the pistils reach their peak of reproductive ripeness. From this point on, the pistils begin to swell and darken slightly, and the tips may begin to curl and turn reddish brown. At this stage the pistillate flower is past its reproductive peak, and it is not likely that it will produce a viable seed if pollinated. Without pollination the calyx begins to swell almost as if it had been fertilized and resin secretion reaches a peak. The pistils eventually wither and turn a reddish or orange brown. By this time, the swollen calyx has accumulated an incredible layer of resin, but secretion has slowed and few fresh terpenes and cannabinoids are being produced. Falling pistils mark the end of the developmental cycle of the individual pistillate calyx. The resins turn opaque and the calyx begins to die.

The biosynthesis of cannabinoids and terpenes parallels the developmental stages of the calyx and associated resin-producing glandular trichomes. Also, the average developmental stage of the accumulated individual calyxes determines the maturational state of the entire floral cluster. Thus, determination of maturational stage and timing of the harvest is based on the average calyx and resin condition, along with general trends in morphology and development of the plant as a whole. The basic morphological characteristics of floral maturation are measured by calyx-to-leaf ratio and internode length within floral clusters. Calyx-to-leaf ratios are highest during the peak floral stage. Later stages are usually characterized by decreased calyx growth and increased leaf growth. Internode length is usually very short between pairs of calyxes in tight dense clusters. At the end of the maturation cycle, if there is still growth, the internode length may increase in response to increased humidity and lowered light conditions. This is most often a sign that the floral clusters are past their reproductive peak; if so, they are preparing for rejuvenation and the possibility of re-growth the following season. At this time nearly all resin secretion has ceased at temperate latitudes (due to low temperatures), but may still continue in equatorial and subtropical areas that have a longer and warmer growing season. Greenhouses have been used in temperate latitudes to simulate tropical environments and extend the period of resin production. It should be remembered that greenhouses also tend to cause a stretched condition in the floral clusters in response to high humidity, high temperatures, lowered light intensity, and restricted air circulation.

Simulation of the native photoperiod of a certain strain is achieved through the use of blackout curtains and supplemental lighting in a greenhouse or indoor environment. The localized light cycle particular to a strain may be estimated from the graph of maturation patterns at various latitudes (p.124). In this way it is possible to reproduce exotic foreign environments to more accurately study Cannabis, Tight clusters of calyxes and leaves are characteristic of ripe outdoor Cannabis. Some strains, however, such as those from Thailand, tend to have longer internodes and appear airy and stretched. This seems to be a genetically controlled adaptation to their native environment. Imported examples from Thailand also have long internodes in the pistillate floral clusters. Thai strains may not develop tight floral clusters even in the most arid and exposed conditions; however, this condition is furthered as rejuvenation begins during autumn days of decreasing photoperiod.

Cannabinoid Biosynthesis

Since resin secretion and associated terpenoid and cannabinoid biosynthesis are at their peak just after the pistils have begun to turn brown but before the calyx stops growing, it seems obvious that floral clusters should be harvested during this time. More subtle variations in terpenoid and cannabinoid levels also take place within this period of maximum resin secretion, and these variations influence the nature of the resin’s psychoactive effect. The cannabinoid ratios characteristic of a strain are primarily determined by genes, but it must be remembered that many environmental factors, such as light, temperature, and humidity, influence the path of a molecule along the cannabinoid biosynthetic pathway. These environmental factors can cause an atypical final cannabinoid profile (cannabinoid levels and ratios). Not all cannabinoid molecules begin their journey through the pathway at the same time, nor do all of them complete the cycle and turn into THC molecules simultaneously. There is no magical way to influence the cannabinoid biosynthesis to favor THC production, but certain factors involved in the growth and maturation of Cannabis do affect final cannabinoid levels, These factors may be controlled to some extent by proper selection of mature floral clusters for harvesting, agricul tural technique, and local environment. In addition to genetic and seasonal influences, the picture is further modified by the fact that each individual calyx goes through the cannabinoid cycle fairly independently and that during peak periods of resin secretion new flowers are produced every day and begin their own cycle. This means that at any given time the ratio of calyx-to-leaf, the average calyx condition, the condition of the resins, and resultant cannabinoid ratios indicate which stage the floral cluster has reached. Since it is difficult for the amateur cultivator to determine the cannabinoid profile of a floral cluster without chromatographic analysis, this discussion will center on the known and theoretical correlations between the external characteristics of calyx and resin and internal cannabinoid profile. A better understanding of these subtle changes in cannabinoid ratios may be gleaned by observing the cannabinoid biosynthesis. Focus on the lower left-hand corner of the chart. Next, follow the chain of reactions until you find the four isomers of THC acid (tetrahydro-cannabinolic acid), toward the right side of the page at the crest of the reaction sequence, and realize that there are several steps in a long series of reactions that precede and follow the formation of THC acids, the major psychoactive cannabinoids. Actually, THC acid and the other necessary cannabinoid acids are not psychoactive until they decarboxylate (lose an acidic carboxyl group [COOHI). It is the cannabinoid acids which move along the biosynthetic pathway, and these acids undergo the strategic reactions that determine the position of any particular cannabinoid molecule along the pathway. After the resins are secreted by the glandular trichome they begin to harden and the cannabinoid acids begin to decarboxylate. Any remaining cannabinoid acids are decarboxylated by heat within a few days after harvesting. Other THC acids with shorter side-chains also occur in certain strains of Cannabis. Several are known to be psychoactive and many more are suspected of psychoactivity. The shorter propyl (three-carb on) and methyl (one-carbon) side-chain homologs (similarly shaped molecules) are shorter acting than pen tyl (five-carbon) THCs and may account for some of the quick, flashy effects noted by some marijuana users. We will focus on the pentyl pathway but it should be noted that the propyl and methyl pathways have homologs at nearly every step along the pentyl pathway and their synthesis is basically identical.

The first step in the pentyl cannabinoid biosynthetic pathway is the combination of olivetolic acid with geranyl pyrophosphate. Both of these molecules are derived from terpenes, and it is readily apparent that the biosynthetic route of the aromatic terpenoids may be a clue to formation of the cannabinoids. The union of these two molecules forms CBG acid (cannabigerolic acid) which is the basic cannabinoid precursor molecule. CBG acid may be converted to CBGM (CBG acid monomethyl ether), or a hydroxyl group (OH) attaches to the geraniol portion of the molecule forming hydroxy-CBG acid. Through the formation of a transition-state molecule, either CBC acid (cannabichromenic acid) or CBD acid (cannabidiolic acid) is formed. CBD acid is the precursor to the THC acids, and, although CBD is only mildly psychoactive by itself, it may act with THC to modify the psychoactive effect of the THC in a sedative way. CBC is also mildly psychoactive and may interact synergistically with THC to alter the psychoactive effect (Turner et al. 1975). Indeed, CBD may suppress the effect of THC and CBC may potentiate the effect of THC, although this has not yet been proven. All of the reactions along the cannabinoid biosynthetic pathway are enzyme-controlled but are affected by environmental conditions.

Conversion of CBD acid to THC acid is the single most important reaction with respect to psychoactivity in the entire pathway and the one about which we know the most. Personal communication with Raphael Mechoulam has centered around the role of ultraviolet light in the bio-synthesis of THC acids and minor cannabinoids. In the laboratory, Mechoulam has converted CBD acid to THC acids by exposing a solution of CBD acid in n-hexane to ultraviolet light of 235-285 nm. for up to 48 hours. This reaction uses atmospheric oxygen molecules (02) and is irreversible; however, the yield of the conversion is only about 15% THC acid, and some of the products formed in the laboratory experiment do not occur in living specimens. Four types of isomers or slight variations of THC acids (THCA) exist. Both Delta1-THCA and Delta6-THCA are naturally occurring isomers of THCA resulting from the positions of the double bond on carbon 1 or carbon 6 of the geraniol portion of the molecule They have approximately the same psychoactive effect; however, Delta1-THC acid is about four times more prevalent than Delta6-THC acid in most strains. Also Alpha and Beta forms of Delta1-THC acid and Delta6-THC acid exist as a result of the juxtaposition of the hydrogen (H) and the carboxyl (COOH) groups on the olivetolic acid portion of the molecule It is suspected that the psychoactivity of the a and ~ forms of the THC acid molecules probably does not vary, but this has not been proven. Subtle differences in psychoactivity not detected in animals by laboratory instruments, but often discussed by marijuana aficionados, could be attributed to additional synergistic effects of the four isomers of THC acid. Total psycho-activity is attributed to the ratios of the primary cannabinoids of CBC, CBD, THC and CBN; the ratios of methyl, propyl, and pentyl homologs of these cannabinoids; and the isomeric variations of each of these cannabinoids. Myriad subtle combinations are sure to exist. Also, terpenoid and other aromatic compounds might suppress or potentiate the effects of THCs.

Environmental conditions influence cannabinoid biosynthesis by modifying enzymatic systems and the resultant potency of Cannabis. High altitude environments are often more arid and exposed to more intense sunlight than lower environments. Recent studies by Mobarak et al. (1978) of Cannabis grown in Afghanistan at 1,300 meters (4,350 feet) elevation show that significantly more propyl cannabinoids are formed than the respective pentyl homo-logs. Other strains from this area of Asia have also exhibited the presence of propyl cannabinoids, but it cannot be discounted that altitude might influence which path of cannabinoid biosynthesis is favored. Aridity favors resin production and total cannabinoid production; however, it is unknown whether arid conditions promote THC production specifically. It is suspected that increased ultraviolet radiation might affect cannabinoid production directly. Ultra-violet light participates in the biosynthesis of THC acids from CBD acids, the conversion of CBC acids to CCY acids, and the conversion of CBD acids to CBS acids. However, it is unknown whether increased ultraviolet light might shift cannabinoid synthesis from pentyl to propyl pathways or influence the production of THC acid or CBC acid instead of CBD acid. The ratio of THC to CBD has been used in chemotype determination by Small and others. The genetically determined inability of certain strains to convert CBD acid to THC acid makes them a member of a fiber chemotype, but if a strain has the genetically determined ability to convert CBD acid to THC acid then it is considered a drug strain. It is also interesting to note that Turner and Hadley (1973) discovered an African strain with a very high THC level and no CBD although there are fair amounts of CBC acid present in the strain. Turner* states that he has seen several strains totally devoid of CBD, but he has never seen a strain totally devoid of THC. Also, many early authors confused CBC with CBD in analyzed samples because of the proximity of their peaks on gas liquid chromatograph (GLC) results. If the biosynthetic pathway needs alteration to include an enzymatically controlled system involving the direct conversion of hydroxy-CBG acid to THC acid through allylic rearrangement of hydroxy-CBG acid and cyclization of the rearranged intermediate to THC acid, as Turner and Hadley (1973) suggest, then CBD acid would be bypassed in the cycle and its absence explained. Another possibility is that, since CBC acid is formed from the same symmetric intermediate that is allylically rearranged before forming CBD acid, CBC acid may be the accumulated intermediate, the reaction may be reversed, and through the symmetric intermediate and the usual allylic rearrangement CBD acid would be formed but directly converted to THC acid by a similar enzyme system to that which reversed the formation of CBC acid. If this happened fast enough no CBD acid would be detected. It is more likely, however, that CBDA in drug strains is converted directly to THCA as soon as it is formed and no CBD builds up. Also Turner, Hemphill, and Mahlberg (1978) found that CBC acid was contained in the tissues of Cannabis but not in the resin secreted by the glandular trichomes.

In any event, these possible deviations from the accepted biosynthetic pathway provide food for thought when trying to decipher the mysteries of Cannabis strains and varieties of psychoactive effect. Returning to the more orthodox version of the cannabinoid biosynthesis, the role of ultraviolet light should be reemphasized. It seems apparent that ultraviolet light, normally supplied in abundance by sunlight, takes part in the conversion of CBD acid to THC acids. Therefore, the lack *Carlton Thrner 1979: personal communication. of ultraviolet light in indoor growing situations could account for the limited psychoactivity of Cannabis grown under artificial lights. Light energy has been collected and utilized by the plant in a long series of reactions resulting in the formation of THC acids. Farther along the pathway begins the formation of degradation products not metabolically produced by the living plant. These cannabinoid acids are formed through the progressive degradation of THC acids to CBN acid (cannabinolic acid) and other cannabinoid acids. The degradation is accomplished primarily by heat and light and is not enzymatically controlled by the plant. CBN is also suspected of synergistic modification of the psychoactivity of the primary cannabinoids, THCs. The cannabinoid balance between CBC, CBD, THC, and CBN is determined by genetics and maturation. THC production is an ongoing process as long as the glandular trichome remains active. Variations in the level of THC in the same trichome as it matures are the result of THC acid being broken down to CBN acid while CBD acid is being converted to THC acid. If the rate of THC biosynthesis exceeds the rate of THC breakdown, the THC level in the trichome rises; if the breakdown rate is faster than the rate of biosynthesis, the THC level drops. Clear or slightly amber transparent resin is a sign that the glandular trichome is still active. As soon as resin secretion begins to slow, the resins will usually polymerize and harden. During the late floral stages the resin tends to darken to a transparent amber color. If it begins to deteriorate, it first turns translucent and then opaque brown or white. Near-freezing temperatures during maturation will often result in opaque white resins. During active secretion, THC acids are constantly being formed from CBD acid and breaking down into CBN acid.

Harvest Timing

With this dynamic picture of the biosynthesis and degradation of THC acids as a frame of reference, the logic behind harvesting at a specific time is easier to understand. The usual aim of timing the moment of harvest is to ensure high THC levels modified by just the proper amounts of CBC, CBD and CBN, along with their propyl homologs, to approximate the desired psychoactive effect. Since THC acids are being broken down into CBN acid at the same time they are being made from CBD acid, it is important to harvest at a time when the production of THC acids is higher than the degradation of THC acids. Every experienced cultivator inspects a number of indicating factors and knows when to harvest the desired type of floral clus ters. Some like to harvest early when most of the pistils are still viable and at the height of reproductive potential. At this time the resins are very aromatic and light; the psychoactive effect is characterized as a light cerebral high (possibly low CBC and CBD, high THC, low CBN). Others harvest as late as possible, desiring a stronger, more resinous marijuana characterized by a more intense body effect and an inhibited cerebral effect (high CBC and CB]), high THC, high CBN). Harvesting and testing several floral clusters every few days over a period of several weeks gives the cultivator a set of samples at all stages of maturation and creates a basis for deciding when to harvest in future seasons. The following is a description of each of the growth phases as to morphology, terpene aroma, and relative psychoactivity. Premature Floral Stage

At this stage floral development is slightly beyond primordial and only a few clusters of immature pistillate flowers appear at the tips of limbs in addition to the primordial pairs along the main stems. By this stage stem diameter within the floral clusters is very nearly maximum. The stems are easily visible between the nodes and form a strong framework to support future floral development. Larger vegetative leaves (5-7 leaflets) predominate and smaller tri-leaflet leaves are beginning to form in the new floral axis. A few narrow, tapered calyxes may be found nestled in the leaflets near the stem tips and the fresh pistils appear as thin, feathery, white filaments stretching to test the surroundings. During this stage the surface of the calyxes is lightly covered with fuzzy, hair-like, non-glandular trichomes, but only a few bulbous and capitate-sessile glandular trichomes have begun to develop. Resin secretion is minimal, as indicated by small resin heads and few if any capitate-stalked, glandular trichomes. There is no drug yield from plants at the premature stage since THC production is low, and there is no economic value other than fiber and leaf. Terpene production starts as the glandular trichomes begin to secrete resin; premature floral clusters have no terpene aromas or tastes. Total cannabinoid production is low but simple cannabinoid phenotypes, based on relative amounts of THC and CBD, may be determined. By the pre-floral stage the plant has akeady established its basic chemotype as a fiber or drug strain. A fiber strain rarely produces more than 2% THC, even under perfect agricultural conditions. This indicates that a strain either produces some varying amount of THC (up to 13%) and little CBD and is termed a drug strain or produces practically no THC and high CBD and is termed a fiber strain, This is genetically controlled. The floral clusters are barely psychoactive at this stage, and most marijuana smokers classify the reaction as more an "effect" than a "high." This most likely results from small amounts of THC as well as trace amounts of CBC and CBD. CBD production begins when the seedling is very small. THC production also begins when the seedling is very small, if the plant originates from a drug strain. However, THC levels rarely exceed 2% until the early floral stage and rarely produce a "high" until the peak floral stage.

Early Floral Stage

Floral clusters begin to form as calyx production increases and internode length decreases. Tri-leaflet leaves are the predominant type and usually appear along the secondary floral stems within the individual clusters. Many pairs of calyxes appear along each secondary floral axis and each pair is subtended by a tri-leaflet leaf. Older pairs of calyxes visible along the primary floral axis during the premature stage now begin to swell, the pistils darken as they lose fertility, and some resin secretion is observed in trichomes along the veins of the calyx. The newly produced calyxes show few if any capitate-stalked trichomes. As a result of low resin production, only a slight terpene aroma and psychoactivity are detectable. The floral clusters are not ready for harvest at this point. Total cannabinoid production has increased markedly over the premature stage but THC levels (still less than 3%) are not high enough to produce more than a subtle effect.

Peak Floral Stage

Elongation growth of the main floral stem ceases at this stage, and floral clusters gain most of their size through the addition of more calyxes along the secondary stems until they cover the primary stem tips in an overlapping spiral. Small reduced mono-leaflet and tri-leaflet leaves subtend each pair of calyxes emerging from secondary stems within the floral clusters. These subtending leaves are correctly referred to as bracts. Outer leaves begin to wilt and turn yellow as the pistillate plant reaches its reproductive peak. In the primordial calyxes the pistils have turned brown; however, all but the oldest of the flowers are fertile and the floral clusters are white with many pairs of ripe pistils. Resin secretion is quite advanced in some of the older infertile calyxes, and the young pistillate calyxes are rapidly producing capitate-stalked glandular trichomes to protect the precious unfertilized ovule. Under wild conditions the pistillate plant would be starting to form seeds and the cycle would be drawing to a close. When Cannabis is grown for sinsemilla floral production, the cycle is interrupted. Pistillate plants remain unfertilized and begin to produce capitate -stalked trichomes and accumulate resins in a last effort to remain viable. Since capitate-stalked trichomes now predominate, resin and THC production increase. The elevated resin heads appear clear, since fresh resin is still being secreted, often being produced in the cellular head of the trichome. At this time THC acid production is at a peak and CBD acid levels remain stable as the molecules are rapidly converted to THC acids, THC acid synthesis has not been active long enough for a high level of CBN acid to build up from the degradation of THC acid by light and heat. Terpene production is also nearing a peak and the floral clusters are beautifully aromatic. Many cultivators prefer to pick some of their strains during this stage in order to produce marijuana with a clear, cerebral, psychoactive effect. It is believed that, in peak floral clusters, the low levels of CBD and CBN allow the high level of THC to act without their sedative effects. Also, little polymerization of resins has occurred, so aromas and tastes are often less resinous and tar like than at later stages. Many strains, if they are harvested in the peak floral stage, lack the completely developed aroma, taste and psychoactive level that appear after curing. Cultivators wait longer for the resins to mature if a different taste and psychoactive effect is desired. This is the point of optimum harvest for some strains, since most additional calyx growth has ceased. However, a subsequent flush of new calyx growth may occur and the plant continue ripening into the late floral stage.

Late Floral Stage

By this stage plants are well past the main reproductive phase and their health has begun to decline. Many of the larger leaves have dropped off, and some of the small inner leaves begin to change color. Autumn colors (purple, orange, yellow, etc.) begin to appear in the older leaves and calyxes at this time; many of the pistils turn brown and begin to fall off. Only the last terminal pistils are still fertile and swollen calyxes predominate. Heavy layers of protec tive resin heads cover the calyxes and associated leaves. Production of additional capitate-stalked glandular trichomes is rare, although some existing trichomes may still be elongating and secreting resins. As the previously secreted resins mature, they change color. The polymerization of small terpene molecules (which make up most of the resin) produces long chains and a more viscous and darker-colored resin. The ripening and darkening of resins follows the peak of psychoactive cannabinoid synthesis and the transparent amber color of mature resin is usually indicative of high THC content. Many cultivators agree that transparent amber resins are a sign of high-quality drug Cannabis and many of the finest strains exhibit this characteristic. Particularly potent Cannabis from California, Hawaii, Thailand, Mexico, and Colombia is often encrusted with transparent amber colored instead of clear resin heads. This is also characteristic of Cannabis from other equatorial, subtropical and temperate zones where the growing season is long enough to accommodate long term resin production and maturation. Many areas of North America and Europe have too short a season to fully mature resins unless a greenhouse is used. Specially acclimatized strains are another possibility. They develop rapidly and begin maturing in time to ripen amber resins while the weather is still warm and dry. The weight yield of floral clusters is usually highest at this point, but strains may begin to grow an excess of leaves in late-stage clusters to catch additional energy from the rapidly diminishing autumn sun. Total resin accumulation is highest at this stage, but the period of maximum resin production has passed. If climatic conditions are harsh, resins and cannabinoids will begin to decompose. As a result, resin yield may appear high even if many of the resin heads are missing or have begun to deteriorate and the overall psychoactivity of the resin has dropped. THC decomposes to CBN in the hot sun and will not remain intact or be replaced after the metabolic processes of the plant have ceased. Since cannabinoids are so sensitive to decomposition by sunlight, the higher psychoactivity of amber resins may be a secondary effect. It may be that the THC is better protected from the sun by amber or opaque resins than by clear resins. Some late maturing strains develop opaque, white resin heads as a result of terpene polymerization and THC decomposition. Opaque resin heads are usually a sign that the floral clusters are over-mature. Late floral clusters exhibit the full potential of resin production, aromatic principles, and psychoactive effect. Complex mixtures of many mon oterpene and sesquiterpene hydrocarbons along with alcohols, ethers, esters, and ketones determine the aroma and flavor of mature Cannabis. The levels of the basic terpenes and their polymerized by-products fluctuate as the resin ripens. The aromas of fresh floral clusters are usually preserved after drying, as by the late floral stage, a high proportion of ripe resins are present on the mature calyxes of the fresh plant. Cannabinoid production favors high THC acid and rising CBN acid content at this stage, since most active biosynthesis has ceased and more THC acid is being broken down into CBN acid than is being produced from CBD acid. CBD acid may accumulate because not enough energy is available to complete its conversion to THC acid. The THC-to-CBD ratio in the harvested floral clusters certainly begins to drop as biosynthesis slows, because THC acid levels decrease as it decom poses, and at the same time CBD acid levels remain or rise intact since CBD does not decompose as rapidly as THC acid. This tends to produce marijuana characterized by more somatic and sedative effects. Some cultivators prefer this to the more cerebral and clear psychoactivity of the peak floral stage.

Senescence or Rejuvenation Stage

After a pistillate plant finishes floral maturation, the production of pistillate calyxes ceases and the plant continues senescence (decline towards death). In unusual situations, however, rejuvenation will begin and the plant will sprout new vegetative growth in preparation for the following season. Senescence is often highlighted by striking color changes in the floral clusters. Leaves, calyxes, and stems display auxiliary pigments ranging in color from yellow through red to deep purple. Eventually a brown shade pre-dominates and death is near. In warm areas, rejuvenation starts as vegetative shoots form within the floral clusters. These shoots are usually made up of unserrated single leaflets separated by thin stems with long internodes. It is as if the plant were reaching for limited winter light. Leaf production is accelerated as plants reach the rejuvenation stage, and resin production completely stopped. Floral clusters left to ripen until the bitter end usually produce inferior marijuana of lowered THC level, especially outdoors in bad weather. Terpene secretion changes along with cannabinoid secretion and psychoactive effect. Various terpenes, terpene polymers, and other aromatic principles are produced and ripen at different times in the development of the plant. If these changes in aromatic principles are directly correlated with changes in cannabinoid production, then harvest selections for cannabinoid level may be possible based on the aroma of the ripening floral clusters. It is important to understand differences in the anatomy of floral clusters for each Cannabis strain. Trends in the relative quantity (dry weight) of various parts (such as leaves, calyxes and trichomes) at various harvest dates are characteristic of particular strains and may vary widely. Some generalizations can be made. In most cases, the percentage of stem weight steadily decreases as the floral cluster matures. Rejuvenation growth can account for a sudden increase in stem percentage. The percentage of inner leaves usually starts very low and climbs rapidly as the floral clus ters mature. This often reflects increased leaf growth near the end of the season. In many strains the percentage of inner leaves drops sharply during the peak floral stage and rises again as calyx production slows and leaf production increases in the late floral stage. Calyx production follows two basic patterns. In one, the percentage of calyxes climbs gradually and levels out during the peak floral stage. It begins to decline in the late floral stage, and leaf production increases as calyx production ceases. Other strains continue to produce calyxes at the expense of leaves, and the calyx percentage increases steadily throughout maturation. In both cases, there is some tendency for calyx percentage to level out during the peak floral stage irrespective of whether leaf growth accelerates or calyx growth continues at a later stage.

Resins generally accumulate steadily while the plant matures, but strains may vary as to the stage of peak resin secretion. Seed percentage increases exponentially with time if the crop is well fertilized, but most samples of drug Cannabis grown domestically are nearly seedless. To determine dry weight, samples are harvested, labeled, and air dried until the central stem of the floral cluster will snap when bent. In plant research, dry weight is done in ovens at higher temperatures, but these higher temperatures would ruin the Cannabis. The dry floral cluster is weighed. The outer leaves, inner leaves, calyxes, seeds, and stems are segregated and each group weighed individually. The percentage is determined by dividing the individual dry weights by the total dry weight. Calyx percentage ranges from 30 to 70% of the dry weight of the seedless floral clusters, depending on variety and harvest date. Inner leaf percentages fluctuate between 15 and 45% of dry weight; stems range from 10 to 30%. It seems obvious that for drug harvesting a maximum calyx production is important to quality resin production. A strain where maximum calyx production occurs simultaneously with peak resin production is a breeding goal not yet attained. Harvesting Cannabis at the proper time requires information on how floral clusters mature and a decision on the part of the cultivator as to what type of floral clusters are desired. With harvesting as with other techniques of cultivation, the path to success is straightened when a definite goal is established. Personal preference is always the ultimate deciding factor.
 
OK. the above article does contain the answer:
On a given flower cluster the calyxes will mature from the bottom up.
Overall the flowers mature at the top of the plant first.

This fits with my own observations. Its a fact I put to good use because after a 'first pressing' where I harvest the top 2/3 of the plant I can come back again. I always harvest straight from the plant because its firm and easy to handle. I leave a few nodes on each stem of buds that I cut. After a few days there will be another a mini harvest from the lower buds and the newly formed buds on the already cut stems. Now the plant has nothing to work on but these few surviving buds and exposed to new light they continue with some vigor. This can go on for weeks or until its not worth getting sticky for. These late nibbles are often the sweetest. :)
 
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