Gene-seed

In Warhammer 40,000, humans are transformed into Space Marines by, among other things, implanting them with a series of 19 organs that dramatically alter and enhance their biology. The organs' DNA strings, from which they are vat-grown, are collectively known as "gene-seed." Go ahead, make your immature jokes.

Origins

Gene-seed is a vital part of the Space Marine program, and is as old as the Astartes themselves. When the God-Emperor of Mankind was still conquering Terra, many of the techno-barbarians' armies had augmented human warriors. The Emperor then realized that he would need vast armies of super-soldiers for his coming Great Crusade, so he embarked on a project to create a mass-producible means of enhancing humans. This project resulted in the twenty Primarchs, and from their genes, he created the gene-seed organs to be implanted into each Space Marine. These organs are usually produced in vats from extracted progenoids but other methods can be used in a pinch, and, of course, the Sinew Coils aren't an actual organ, so they can be produced much more readily, like any other cybernetic implant.

The traditional loyalist method of implanting gene-seed into recruits until it grows into new organs, then retrieving fresh gene-seed from them happens to loosely resemble the real-life concept of adult stem cells, invaluable for life-saving surgeries and keeping your liver from shriveling up when you get too drunk.

The process was originally available on a nearly industrial scale for the Great Crusade - both Terra and each legion recruitment worlds could make up the numbers for thousands of recruits. Occasionally they could be inducted as adults, although the limitations of this varied and usually an adult recruit was worthy but couldn't take a gene seed. In this case various surgeries and augmentations could turn them into a 'demi-marine', although the difference in performance was notable. A notable exception was Magnus the Red's teacher Amon, who would have been far too old for geneseed implantation but was altered by Magnus' powers to accept the implants.

During the Horus Heresy the Traitor Legions came up with a quick and dirty method for boosting their ranks by taking hundreds of thousands of aspirants (any sufficiently tough psychopath among their millions of followers) and forcing them through the implantation process much more rapidly than normal, along with substituting the years long training process with constant hypnotherapy training. The success rate was small but they had so many potential recruits it still resulted in a boost in their numbers. These Newborn, as they came to be known, were equipped with the same gear as regular marines but utterly lacked the skills and prowess of a traditional legionary. Their forced creation also led them to being seen as second-class citizens amongst their legions, and thus they created their own subcultures and traditions. As a result, they ended up being used as disposable shock troops whilst regular Astartes were saved for more important duties.

The Loyalists guarding Terra also sped up their training processes before the Siege, although they were more discerning with their recruits and still provided hands-on training. Notably, these recruits frequently did not come from Terra or their Legion home worlds, but from the worlds bordering the Sol System. As such they occupied an odd place in the legions social structure.

After the Heresy, the loyalist legions and then chapters scaled back the process, relocating all the materiel to their homeworlds or fleet apothecarions. As a preventative measure learned from the Heresy, subsequent recruits were taken at a younger age, both to screen out weakness as soon as possible, and to more readily indoctrinate the prospective marine to being loyal (YMMV). Most chapters also introduced far more stringent levels of hypnotherapy and indoctrination than any Heresy-era Marine would ever have had, and their standards for general recruits were also much higher, since now they recruited fewer aspirants from a smaller population (relatively) instead of a lot from a massive group, which could even be from several different worlds in cases of emergency recruitment.

The Organs

Oddly, most 40K fluff never refers to most of these organs, especially when situations come up where these organs might have been especially useful. The worst offenders are usually the sensory ones; the Occulobe, Lyman's Ear, and Neuroglottis, taken together, ought to make them notice things normal humans can't, especially under low-light conditions, but the fluff usually has them only picking up the standard level of detail.

1. The Secondary Heart is pretty self-explanatory. It lets a Space Marine get his blood flowing at a faster rate through his body and probably to the more critical bits, which lets his muscles work harder and longer, and it gives him a backup just in case one fails (i.e. gets shot, stabbed, ripped out by Guilliman, or what have you). The Secondary Heart is not beating at all times. It only activates during times of stress (bear in mind that a twenty mile run is not considered stressful to an Astartes); otherwise, it just sits there awaiting activation.
2. The Ossmodula makes a Space Marine's skeleton grow and incorporate some ceramic dietary supplements into itself and fuses his ribs together into a solid structure. The Tyrant guard's fused ribcage hints that the Hive mind has incorporated Space marine DNA into-*BLAM* convergent evolution is at work. Ironically, the fusing of the rib cage would be considered a bad thing, as the Space Marine would not be able to breathe properly-, as pointed out by Fabius Bile the ribs are not one solid plate but actually many smaller plates with near invisibly small spaces between that allow them to flex and achieve a proper range of motion (so similar to Laminar armor instead of full Plate). Note another explanation is the Multi lung (see bellow) may help push the ribs apart harder than a normal pair of lungs ever could. Also a fully fused rib cage would cause a loss of flexibility and may be an undocumented Ultramarines mutation since the vast majority of them walk around like they've got a broom handle shoved up their asses.
3. The Biscopea triggers muscle growth. These first three organs are often implanted together, as they collectively prepare the Initiate's body for further augmentation. Around this time, the Initiate gets put on a diet containing extra nutrients to fuel his new growth.
4. The Haemastamen enhances a Space Marine's blood to better accommodate the Space Marine's nifty set of new organs by improving its ability to carry oxygen. It also turns it bright red.
5. The Larraman's Organ creates Larraman's Cells, which are like super-platelets: they make wounds rapidly scab over. This allows Space Marines to keep fighting even when shock and blood loss would kill most people.
6. The Catalepsean Node messes with the body's sleep cycle, letting Space Marines get away with four hours of sleep every day, or even less, and making it possible for them to not lose awareness, even after like 2 weeks without sleep. It also makes the brain susceptible to hypno-therapy, which is a fancy word for "brainwashing" the Initiates while they are asleep. As a nasty side effect, it allows Chaos marines to re-brainwash loyalists they captured alive (or better: in suspended animation). Unsurprisingly to anyone familiar with Chaos, this trick doesn't work in reverse, which might just go to show that the Chaos Marines might be closer to the true human nature than the loyal Marines... or more likely that Chaos is corrupting as fuck.
7. The Preomnor is like a bird's crop, in that it stores food before passing it to the stomach, except that the Preomnor also decontaminates its contents as well. In Ian Watson's Space Marine novel, Initiates celebrated getting this organ by eating the most disgusting things they could until they finally provoked a vomit reflex.
8. The Omophagea is one of the odder organs. It connects to the stomach and the spinal cord, and somehow takes knowledge from the brain tissue that a Space Marine can eat. Interestingly, most Space Marines are pretty reluctant to actually eat brains, as the process is physically extremely unpleasant, leaves them in a vulnerable torpor whilst they process the information, and is one of the very few things that can readily make a Space Marine vomit afterwards. The Thousand Sons (pre-heresy) - contrary to their civilised, haughty persona - highly approved of the practice. And, in fact, it is not just the brain, since some Astartes rituals, if not chapter names, originated from this; the Blood Angels and their descendants are especially famous for blood drinking and rip and tear in general; this is actually because they receive knowledge and visions from it. What? You thought the Blood Angels had visions of Sanguinius only to have a grimdark, mystical aura? Before reuniting with their Primarch, the Blood Angels standard operating procedure was to ritually cannibalize their dead officers and sometimes even assume their rank and identity. Additionally, the memories they absorb become very hard to forget (brain washing technologies can help). Now just add some chaos to that mix and you've got heresy brewing...
9. The Multi-Lung... do I have to spell it out? It's an additional lung! It also has a lot of filters and the like, and can function underwater and in low-oxygen atmospheres. In other words, it allows the Space Marine without a helmet to breathe under water and in toxic air. Just how does this third lung fit in with the other two Marines are born with and all the other organs implanted later? Very carefully, that's how.
10. The Occulobe gives Space Marines super-vision, and allows their eyes to adjust to changes in light almost instantly, or at least in the fluff.
11. The Lyman's Ear gives them super-hearing and makes them immune to motion sickness and concussion. However, in-game, they have no particular immunity or even resistance to concussion, probably because it's often delivered by a hammer to the head.
12. The Sus-an Membrane lets them enter a state of Suspended animation when they are critically hurt or starving. This serves the purpose of conserving a mortally wounded marine until his body is recovered and healed by apothecaries. So each time you remove a marine model on the tabletop, chances are (fluff-wise) it has entered sus-an, rather than died. In fact, on most campaigns that's what happens to your characters. It should be noted, marines cannot leave suspended animation state at will, and can only be "awakened" through the use of specific drugs.
13. The Melanochrome implants into the lymph nodes, and makes Space Marine skin like transition lenses: it darkens in response to radiation, rapidly producing reinforced melanin that protects against both UV and gamma. On the plus side, they tan really fast, but the fact that they pale right back up once they're inside armour (unless they are Salamanders) is not that great.
14. The Oolitic Kidney detoxifies their blood, like their regular kidneys, but this one does it really fast. Also, it controls some of the other organs' hormones. As a result, Space Marines are unaffected by most poisons short of what the Dark Eldar or Tyranids can produce. It also means they can't get drunk, though the Space Wolves found a way around that by creating Mjöd, an extremely toxic drink made of local plants of Fenris, that temporarily neutralizes the Oolitic kidney's ability to filter toxins. Another organ which ceases to work on the tabletop, as Space Marines have absolutely identical reactions to Poison as ratlings, ogryn, and standard humans. Of course, considering that the poisons you find on the tabletop are Tyranid, Dark Eldar, and Nurgelitle shit, we can probably write this one off. Also, in some fluff, it saves the bearer from poison by causing them to enter a sort of super-metabolic coma during which they burn off any toxins while unconscious and feverish—so, much like with the sus-an membrane, it allows you to say that any of Your Dudes taken out by Poison effects are just temporarily disabled.
15. The Neuroglottis implants into the upper nasal passage, drastically improving both smell and taste; lets a Space Marine identify chemicals and track adversaries by taste. Blood-hound much? Apparently, the Space Wolf version of this is so good that they designed a special Wolf Helm that lets them use their sense of smell even when fully armored, and is also why they otherwise avoid helmets as much as possible (and definitely not just so that they can gel their hair into giant crests and/or grow chest-length Santa beards).
16. The Mucranoid gives Space Marines super-sweat that protects them from heat and cold, and can even be hardened into a vacuum-proof shell. Remember that awesome scene where Guilliman boxes Word Bearers in space? Yep, he did it while covered in that thing.
17. The Betcher's Gland gives them acid spit, which is probably the only bonus for going to combat without a helmet. The acid within varies depending on the source, from simply reacting only to organic tissue to fully dissolving ceramite. Some Chapters, such as the Blood Angels and the Iron Snakes, consider using the Betcher's Gland in combat to be dishonourable. Technically, the acid spit is just a fringe benefit of the gland’s true purpose—allowing marines to chew through everything from tough xenos carcasses to prison bars. In any event, Space Marines on the table-top do not have access to acidic spit, unlike e.g. Tyranids. (Or maybe they do; space marine ancients get a lot of attacks for someone with their hands full.)
18. The Progenoids provide no benefit to the Space Marine, but are universally considered the most important organs, as they are the keys to making more of the organs above. Each Space Marine starts with two, one in the neck and one in the chest, and they are removed after their death. The neck-based progenoid is ready for harvesting five years after implantation, while the chest-based progenoid is ready after ten. Each harvested progenoid is a bunch of DNA and chemical stuff, that, when implanted in a suitable environment, grows into more organs. Chapters that took severe losses and could not reclaim the gene-seed of their brothers basically take random people with an affinity to the Astartes gene-seed (or vat-grown bodies similar to the ones they use for Servitors), implant them with all space marine organs, wait ten years, and slaughter them to harvest the gene-seed. Sometimes, the term "gene-seed" is taken to refer to these glands specifically.
19. The Black Carapace interfaces a Space Marine with his armour, making them effectively one being. It also serves as back-up armour, providing a level of protection similar to a flak jacket, in case a marine somehow ends up running around naked. Space Marine Scouts don't have this yet, because of the Codex Astartes, and it's noted as the only one of the original organs that has to be implanted whole rather than potentially grown inside the neophyte as a germ-seed (a la Insanguination). They don't get it (and thus an expensive suit of power armor) until they've spent a decade or more in a Scout Company. Tyranids nicked this for their tyrant guard, which makes you think how did the foot-slogging hive tyrants survive anything beforehand. As clearly established by how humans perform in Power Armor, this organ does nothing on the table-top; Sisters of Battle, for example, get just as much (or more - they can lift a lascannon by themselves, unlike humans in mere carapace armour) benefit from Power Armor as Space Marines do. The Deathwatch RPG explained it as allowing a “Size (Huge)” Space Marine in Power Armor to not suffer any movement penalties—which does seem impressive if you’ve ever seen someone in craft foam Astartes armor trying to hobble around.
20. The Canis Helix is not an organ, but considered nonetheless to be part of the gene-seed; it appears to be a genetic cocktail brewed by the Wolf Priests of the Space Wolves, and while it induces a level of mutation beyond that seen in any other chapter, Russ' gene-seed organs appear to have a 100% rejection rate in humans who do not first take the cocktail.

Primaris Organs

After Roboute Guilliman told Belisarius Cawl to make a next generation Space Marine, he ended up creating Marines that had an enhanced version of the Astartes Gene-Seed, which contained an extra three organs (possibly due to taking elements of the Custodes or even the Primarch creation process). While some have expressed concern of the project, some due to worries about how some of the mutations of some of Chapters with more exotic mutations and some due to the belief that the Emperor's work shouldn't be tampered with, Cawl claims that the Primaris Gene-seed is more stable than the regular Astartes gene-seed, with an estimated deviancy of .001% per generation. How true this claim might be is up for debate, but Cawl has created Space Wolves successor chapters that haven't immediately turned into Wulfen so it may have some truth in it. It's also worth noting that Cawl did something to make the Primaris Marines physically incapable of turning traitor. Guilliman's internal monologue in Dark Imperium reveals that he considers it an impossibility for Primarines to turn, unlike "the old breed". This has interesting implications about whether or not the Primaris marines truly have free will.

As of recent fluff updates/leaks, the new organs are as follows.

1. The Sinew Coils essentially modify the muscle strands and makeup of the Primaris Marines, buffing their pecs and biceps to the 10th degree. Whilst one would assume it would resemble something from Inspector Gadget's coil legs, the truth is....it is pretty much like that in function. These are reinforced coil-cables made of durable supermetals that are capable of contracting with incredible strength, making Primaris stronger and tougher than others. Even outside of their power armor, swinging a sword at a Primaris might just bounce off his hot and sweaty muscles because of these coil cables, and it amplifies his strength to the point where he can crush a human skull with his bare hands or bite through metal cable like it were jerky.
2. The Magnificat may sound a bit lame, but its function is actually what makes the Primaris 'bigger, better, faster and stronger' then regular Astartes. It produces a mix of hormones that greatly enhance the effects of all of the other genetically engineered organs that float around in the Primaris. So every Primaris gets more bang for his genetic buck. Intriguingly, it's stated to be only half of another super-organ dubbed the Immortis Gland originally made for the Primarchs, but Cawl was only able to find the materials and genetic blueprints needed to make the right half; the information needed to make the left half had seemingly been erased. The function of the complete Immortis Gland is unknown, but given that its other name is "The God-Maker" and the fact that only the Primarchs are known to have the organ, it stands to follow that it is likely what amplifies their abilities to the level of demigods.
3. The Belisarian Furnace is an organ that is named after the Galaxy's least humble Archmagos. This one is a last-chance fail-safe kind of thing. The Belisarian Furnace is an organ that activates when a Primaris is on death’s door, giving them a great burst of super-adrenaline (not the actual term) and boosted regeneration, but helping to keep them alive even at great cost, so that a Primaris can take out as many of the enemy as possible even as his flesh and bone rapidly regrow themselves. This allows fan-fic writers to create Hollywood-esque 'final pushes' without making it look stupid, which should make for some amazing last stands. It also appears to be vital for the Rubicon Primaris procedure intended to convert existing Space Marines into Primaris Space Marines, as the procedure temporarily kills the subject in its current form. The organ doesn't just activate during fights however ; the fluff has shown a Primaris actively trying to "calm down" and stop their Furnace activating whilst they were critically injured whilst they were being rescued as the resulting physical shock would finish them off.

Implantation

Not everyone is compatible with implantation; potential candidates are screened for tissue compatibility, and kept young, as being post-pubescent has a marked increase in the tissue rejection rate (and being actually elderly makes it even worse); even so, not all candidates survive the implantation process. The Great Crusade era had inductees from a variety of ages, possibly because there were more resources, candidates, and understanding of the tech to compensate for failure rates. Back then a one-in-two success after screening was considered pretty good. These days the screening is insanely brutal amongst Loyalist chapters, with most aspirants dying before being even considered for implantation, but the success rate is accordingly much higher. Youths are also still preferred post-Heresy as they are easier to mould and condition into loyal Marines than teens or young adults.

One of the most obvious limits of their design is that the Emperor never solved the problem of making them compatible with female hosts (although some argue that this was deliberate as a way of ensuring that Space Marines could not breed true, nobody can be entirely sure), so the very first screening step is usually by sex. Furthermore, not all organs can be implanted at once, as some prepare the body for others, and the body needs time to adapt to and accept the implants. The maximum speed process has been described in the fluff as taking around 6 years and assuming the soon-to-be Marine will be a Primaris is:

1. If you are the Space Wolves, the Canis Helix.
2. Secondary Heart, Ossmodula (bone growth), and Biscopea (muscle growth).
3. Sinew Coils (tendon/ligament reinforcement), Magnificat (enhances all other geneseed organs), and Belisarean Furnace (combat stimms, attaches to hearts).
4. Haemastamen (improved red blood cells) and Larraman's Organ (improved platelets).
5. Catalepsean Node (brain implant: sleep like a dolphin).
   • If this is an actual Space Marine, begin hypnotherapy/brainwashing only after this implant.
6. Preomnor (gullet), Omophagea (flatworm stomach), and Multi-lung (third lung).
7. Occulobe (improved visual cortex).
8. Lyman's Ear (replacement outer, middle, and inner ear).
9. Sus-an Membrane (brain implant: self-induce coma).
10. Melanochrome (lymph implant for radiation resistance).
11. Oolitic Kidney (enhanced liver/kidney) and Neuroglottis (improved inner nose).
12. Mucranoid (secretable space suit).
13. Betcher's Gland (extremely acidic saliva).
14. Both Progenoids (for reproduction).
15. Black Carapace (for improved power armour compatibility).

The largest time gap between implants is between the Betcher Glands and the two last ones on the list, and by that time the Marine already serves as a Scout. Once he is deemed physically and mentally ready, he will recieve the Progenoids and the Black Carapace that turns him into a full-fledged (Primaris) Marine.

The Rubicon Primaris most likely entails the implantation of the three Primaris gene-seed organs past the normal time that gene-seed would normally be implanted, though the specific order of implanting in this scenario is currently unknown, however it is known that the Furnace resuscitation is needed to finish the process so it might come in first. The logical follow up is the enhancing Magnificat and finished with the Sinew Coils, the pain of which would cause clinical death through shock, brute forcing the Furnace into activating and charging the Marine up.

Tabletop Effect

Most of the organs have no visible effect in the game, at least if the in-game rules are an indication; here are how the organs seemingly manifest (or not). For some reason, Space Marines aren't any faster at running than an unaugmented human and are slower than fast humans, such as Sisters of Silence. Most likely, this lack of impact is either for balance purposes or because it can't be properly represented using in-game means.

• Has an effect, at least in aggregate:
  • Secondary Heart, Ossmodula, Biscopea, Haemastamen, Larraman's Organ, Multi-Lung, Oolitic Kidney, Mucranoid: +1 Strength and Toughness, relative to a normal human.
  • Sinew Coils, Magnificat, Belisarean Furnace: +1 Attack, can only be wounded on a 4+ (1CP strat activation).
    • Was also +1 Wound in 8th edition. As of 9th edition the Firstborn also get +1 Wound.
  • Catalepsean Node + Hypnotherapy: And They Shall Know No Fear, +1 Ld.
  • Progenoids: These are given as the reason an Apothecary loses his turn if he fails a resurrection roll, as he is busy collecting them.
• Effects limited to fluff:
  • Occulobe: Mostly ignored; with or without his helmet on, most Space Marine's vision is never considered any better than an unaugmented humans, even when distance or lack of light are explicitly causing vision problems. It may however be a reason why some Space Wolves get the 'acute senses' rule.
  • Lyman's Ear: Demonstrably ignored under the 8th edition rules; Space Marines behave exactly like unaugmented humans when exposed to sonic weaponry, when needing to notice things they can't see, or when in need of balance. In previous editions, this would presumably have contributed to a higher Initiative stat.
  • Neuroglottis: As noted above with the Lyman's Ear, as this would predominantly serve to supplement that, given some tabletop rule for Space Marines actually having improved senses which only the Space Wolves have.
  • Melanochrome: Demonstrably ignored; Space Marines respond to photonic radiation (usually provided by the Adeptus Mechanicus) exactly like unaugmented humans do.
  • Black Carapace: Demonstrably ignored; many human factions have access to Power Armour, particularly Sisters of Battle, and none have ever been shown to be even slightly more encumbered by Power Armour than Space Marines are.
  • Preomnor, Omophagea, Sus-an Membrane, Mucranoid, Betcher's Gland: These have no in-game effect, but would typically not be expected to, as a Space Marine with his suit on and participating in a battle would not have call to use them, although the Betcher's Gland could be implemented as a very, very short range weapon, in principle.
• Alternate Take: The Occulobe, Lyman's Ear, and Neuroglottis could grant the +1 Initiative marines enjoy(ed prior to 8th). The Melanchrome also likely adds to toughness.

Gene-seed vs. Genetic Engineering

A lot of people (including, it must be admitted, Games Workshop itself) often think of gene-seed as a genetic modification program, but this is inaccurate. Gene-seed does fall under the blanket of "bio-enhancement" or "bio-engineering"—that is, the augmentation of the physical form. However, gene-seed is based on the physical implantation of artificially designed and engineered organs, whereas genetic engineering would modify the DNA of the aspirants to naturally develop the various traits. In essence, Space Marines are what Rifts would call "bio-borgs": organisms biologically modified by artificial biological implants. Fleshy cyborgs, essentially. This is an important difference, as it allows the Space Marines to wriggle around that pesky "thou shalt not modify the holy human form" law of the Imperium. From a genetic standpoint, a Space Marine is probably closer to human than most abhumans are, and so they fulfill the letter of the law (if not its spirit - but hey, the Emperor made the rules, so he can break them as much as he wants, 'cause he's the motherfucking Emperor). It also has the side effect of making sure that Astartes can't breed true-ish (assuming they can breed, a point GW can't ever make up their minds on, any children they had would be more likely compatible for geneseed then a baseline human but by themselves would not be space marines, just more likely to be able to be), and thus will always have to rely on baseline humanity rather than replace it, just in case they start getting too big for their transhuman britches.

• Note: Like Mitochondria (the powerhouse of the cell), the implanted organs carry their own DNA, cloned from the Primarchs, thus turning the marines into chimeric humans. So yes indeed, bio-borgs, but ones that show characteristics of DNA foreign to the host's body. Example: is there a specific organ that gave Sigismund "a patrician face that echoed the same austere lines as his liege lord, Rogal Dorn"? Let us be reminded, Sig was terran, and Dorn grew up on another planet (he was made by the Emperor in a Petri dish, does that count as being born on Terra too?). Who knows what he looked like when he was mortal, he now looks like a son of Dorn. In fact, most Astartes end up looking similar to their parent Primarchs, like Horus "little Horus" Aximand looking like Horus "fucking Horus" Lupercal. Of course, this is because GW thinks it is bioengineering, but at least it's "Human Chimeras" instead of "Humans with furry creatures", that'd be Heresy, right Emps?
  • RIGHT
  • As it happens, for Sigismund, the answer is yes—the Ossmodula would alter the shape of his skull, and the Biscopea the arrangement of his facial muscles—but the fact remains that GW routinely treats gene-seed as if it were gene therapy, rather than implants.
  • This cloned DNA is also why every body freaks out over gene seed stocks from the fallen primarchs in case their geneseed has some flaw that increased its susceptibility to chaos.

The Tithe and New Foundings

As part of the deal in Guilliman's Codex Astartes, the Astartes were afforded almost complete autonomy from the Administratum, which means they don't have to pay Imperial Taxes except for a tithe of 5% of their gene-seed which they must send to Mars.

This has a few important purposes:

1. To monitor the genetic purity of the Space Marine Chapter as a whole and make certain a chapter is not suffering heretical mutations.
2. To create new foundings of Space Marines whenever the High Lords of Terra deem it appropriate to do so.
3. To act as an emergency reserve of gene-seed in the unlikely event that the Chapter's existing gene-seed stocks are lost or tainted.

Note that even though the Adeptus Mechanicum will have stockpiles of a chapter's genetic material, it is not frequently used as a contingency for a space marine chapter if things get a bit hairy. Several times in the fluff a Chapter has faced extinction due to high losses (see Black Consuls, Celestial Lions, Lamenters, Marines Errant, Scythes of the Emperor yadda yadda yadda) but they'd never thought to ask Mars for their gene-seed back. The only known instance of it happening is in the 8th Edition Death Guard Codex, where the Minotaurs were forced to undergo a mission to Mars when a Death Guard assault on their fortress-monastery resulted in the poisoning of their gene-seed stocks, but the Minotuars are basically the High Lords personal beat stick against other marines so perhaps this is special treatment.

It's also a bit uncertain how chapters with particularly negative defects manage to keep their genetic condition concealed, if the government has access to their core materials. Some chapters get around it by sending only the "purest" gene-seed to Mars and so come out of the inspection smelling of roses, but when the condition is widespread (as it is with the Blood Angels) it probably only highlights how little the Mechanicus actually understand about the Emperor's work. (Things might perhaps change now that Cawl is back, though.)

It's also important to note that each chapter's gene-seed is kept in isolation. i.e: Just because a chapter claims descent from the Ultramarines does not actually make their gene-seed the same as their forefather. It's basic evolutionary biology that when a population splits and develops elsewhere it starts forming its own traits advantageous to its circumstances and environment. Hence the Mortifactors gene-seed will largely be very different from the Howling Griffons, despite sharing a common ancestor. So rather than being genetic "brothers" as they would be within a chapter, different Space Marine chapters sharing a lineage would be more like nephews or cousins.

When the time comes around to found new chapters, the adepts will look for favourable traits that they want to replicate, although the Ultramarines are cited as being the most utilised source for new foundings, thus any new chapter of Ultramarine descendants would likely be second generation Ultramarines and be quite close to their forefather genetically at least until the chapter adapted to its founding homeworld or particular circumstances. But that doesn't prevent those descendants themselves from being used as templates in later foundings, so after several generations you could end up with something completely different from the original baseline stock.

Mutation

As each new set of gene-seed comes from a Space Marine, requires many years to mature, and many more to be used, there is a lot of room for things to go wrong. This is especially true when desperate circumstances (the impending Great Crusade, or a Chapter being dangerously under-strength) cause Space Marines to skimp on the safety checks and wait periods in order to accelerate Chapter growth. Over the millennia, many flaws have found their way into into the gene-seeds of each Legion, and some of their successors have even more - mutations typically get worse as you make copies of copies. Further mutations have been introduced, of course, in foundings featuring heavy tampering by the Adeptus Mechanicus (see the discussion of the 21st founding, below). Also one should bear in mind that Legions and Primarchs were artificially engineered to fill different niches, so some of these might be not acquired mutations, but deliberately added quirks to better fit in their roles.

No Known Mutations

Dark Angels (I Legion)

No defects, unless you consider their rampant paranoia and secrecy STAUNCH LOYALTY AND HONOUR as such. Ties with Ultramarines for "most stable source", so the limited number of Dark Angels successors is usually put down as the Inquisition's limited knowledge of the Fallen Angels. That and the fact that due to successor chapters often working with/take orders from the Dark Angels has lead to the Inquisition to fear the Angels and successors to be "legion building" THE MOST LOYAL CHAPTERS DUE TO THE PURITY OF THEIR GENE SEED. Although that would be more credible if they had ever been anything less than a Legion.

Iron Hands (X Legion)

• No defects.
• Noted predilection towards machinery and a disdain towards flesh.

And just to make sure, if there are any, they follow the Bible's advice on offending organs and replace them with holy machinery instead. Would probably be a very desirable source for new chapters if not for the absolute devastation they suffered in the Drop Site Massacre leaving them barely recovered millennia later and needing all the gene-seed they can get for themselves. Because, you know, even though there has been more than enough time for more than twenty other foundings of whole, brand new chapters, and some of those foundings likely making new foundings of their own, the Iron Hands still haven't had enough time. It also doesn't help how they tend to be total pricks at times. Or to have internal disagreements on policy so hard 'resolving' them had Chapter-splitting consequences. which does not help building up Marines/gene-seed reserves, and gets you overlooked even more when the Mechanicus is deciding on whose reserves to use for the next Founding.

Alternate take: You need causalities that don't destroy the body entirely to harvest geneseed, or you need to implant it into clones, both of which are unlikely for a chapter that uses augmentations with power supplies that explode like melta bomb and that abhors even the organic parts of their own servitors. No new gene-seed, no new mutations.

World Eaters (XII Legion)

I know what you're thinking, but it's caused by an additional cybernetic implant - the Butcher's Nails - rather than anything in the gene-seed itself. While not technically part of their gene-seed, it has been carried on as a tradition by all the "successors" that resulted from the Legion's fracturing after the Battle of Skalathrax. Hell, they were known to be total bros before all the trouble with nails. And now they are little more than Teamkilling Fucktards.

Khorne might have given them one blessing though, a sort of extra lives cheat code… But they are limited and are only earned by participating in the arenas.

Ultramarines (XIII Legion)

• No official defects. Considered the most stable and preferred source of gene-seed for new Foundings. Possible predilection towards conformism and communitarianism.
  • Due to a lack of any real interesting or identifying traits in and of itself, discussions for reclassifying the Ultramarine gene-seed from the most stable source to the most generic source were in the works. These talks were surreptitiously shut down by Inquisitor Warden for offending his sensibilities.
  • Also worth noting is that the Ultramarines, in one of their bro-iest moves, "adopted" a fair number of loyalists from traitor legions after the heresy and are heavily implied to have taken in members of the two missing legions after whatever it was that got them disbanded. There is no real way of telling if an Ultramarines "successor" might be from the relatively stable stock of World Eaters or Iron Warriors. In most cases only a single high-ranking archmagos knows the truth, so even the chapter itself (except possibly the command and the eldest brothers) is in the dark about its history.

Death Guard (XIV Legion)

• Emphasis here is on known mutations - there's no actual evidence of Death Guard gene-seed having any problems at all, because we've never gotten any fluff covering any attempts to use the gene-seed from a Death Guard, even though one would assume that by now they'd have the most fucked up Progenoids of any Legion.

During the Great Crusade, the Death Guard had a much stronger constitution than most Legions, leading to their role as the toxic environment specialists. Post-heresy, however, the Death Guard are so diseased that implanting Aspirants seems more or less impossible - however, there have been new Death Guard legionaries who are not defectors from other Legions or renegade Chapters, apparently made with the gene-seed reclaimed by the Plague Surgeons. How it functions still, no one knows, though we suspect it has more to do with Nurgle infectiousness than with the progenoids themselves.

The most overwhelmingly likely explanation is, however, the Death Guard simply jacking loyalist gene-seed from fallen loyalist astartes and loyalist gene-seed stockpiles and using it to create new recruits, as every single Traitor Legion with tainted, unusable geneseed does this. As per LoS, appears to be a mixture of both, likely depending individually on each Death Guard members level of disease and mutation, supplemented by stolen and possibly altered loyalist gene-seed (see Plague Surgeon).

Alpha Legion (XX Legion)

No such Legion exists, therefore no such gene-seed exists.

• Were you really expecting known mutations, or anything else, from this Legion???

In all seriousness; like much of the Alpha Legion, we don't know all that much about their gene-seed. Whatever idiosyncrasies or flaws they have are kept under very tight lock and key. And with their nature of running as individual operatives in and out of the Eye of Terror, to what degree their gene-seed is tainted by the Warp or other means varies from Marine to Marine. Though their knack for secrecy, paranoia, and tendency to subvert the enemy instead of direct combat is probably a notable trait. Outside of occasional Warp-corruption, we assume it's possibly one of the most stable gene-seeds, probably as much as the Imperial Fists or Night Lords. Something of note; they often use any mutations that they have as a scare tactic, hiding them and then revealing them on the battlefield in order to shock the opponent. They also use hypnosis and deep-indoctrination more than other chapters, so they may have an altered sus-an and/or catelapsean. Sometimes it's implied they all look alike (like Alpharius) but sometimes this is said to be due to surgical alterations, but it could be that the gene-seed does more to alter their appearance than normal, like the Sons of Horus and this is further supplemented by surgery. It's also said that some Alpha Legionnaires tended to be noticeably taller than most Space Marines, large enough to pose as either Alpharius or Omegon. This, combined with their unusually short Primarchs and their tendency to look alike, seems to suggest that this was intentional, either by the Legion themselves picking only notably large aspirants or by the Emperor fully leaning into the Legion's whole "smoke and mirrors" theme.

Minor Mutations

These chapters may have cosmetic or easily-fixed mutations.

Emperor's Children (III Legion)

• Implicitly defective and/or enhanced Lyman's Ear, and possibly Occulobe and Neuroglottis.
  • An accident (non Chaos or other enemy action related) during the early days of the raising the Legions resulted in only two hundred Space Marines of the III Legion by the time the Emperor found Fulgrim and Chemos, forcing the Emperor's Children to work alongside the Luna Wolves for the early decades of the Great Crusade; due to Fulgrim's high standards of perfection being passed on the Legion's apothecaries, Emperor's Children Aspirants had to be nothing less than physically perfect, resulting a largely stable gene-seed, including forbidding psykers to join the ranks. By the 41st millennium, however, the Emperor's Children's gene-seed is fucked up beyond all reason. Millennia of drugs from both inside and outside the Warp have damaged it greatly. While the relatively few sane tech-priests left and Dark Mechanicus working with them can and do make new Marines (thus proving their Marines can and do make new gene-seed), the chemical and other material abuse render each generation even crazier. Even before the Heresy, they were more sensitive to external stimuli, like sound, implying a defective Lyman's Ear with reduced ability to filter out sounds the Marine doesn't want to hear, and potentially similarly defective Occulobe and Neuroglottis.
• Initial batches of gene-seed were defective, carrying an infectious a super-cancer/progeria creatively called "The Blight" it also caused the entire legion's gene-seed stock on Terra to rot into useless sludge. Thankfully after Fulgrim's return the rotten batches were discontinued and replaced with fresh clean gene-seed preventing it from becoming a permanent feature of the legion. The Blight may be more of a spiritual infection because even after dying and having his soul swapped into new bodies Fabius Bile still suffers from all the symptoms.

Iron Warriors (IV Legion)

• Implicitly defective Ossmodula
• Implicitly enhanced Progenoids:
  • They have the lowest gene-seed rejection rate, which enables them to replenish their huge number of casualties with an even higher number of recruits.
  • Minor Ossmodula mutation sometimes causes limbs to twist and deform as the marine grows really old (we're talking hundreds of years to millennia) - this is usually dealt with by replacing limbs with bionics, or cutting them off and placing the cripple into a Dreadnought, Helbrute, or some weird custom made dreadnought-esque walker, since due to their specialty and organization only high-ranking officers can survive for so long, and they surely ain't going to suffer from the "Crazed" rule.
  • Below average resistance rate to Bionics.

Especially noteworthy for the Iron Warriors considering they reside in the Eye Of Terror, there has been little mutation in their gene-seed as well. Those that do suffer mutation simply replace their mutated limbs with cybernetics. However, there may be some unknown issue which caused them to be afflicted with the Obliterator Virus (or was it they who made it? It's been confirmed Perturabo and Mortarion worked together on the obliterator virus) More easily - the first ones originated from them.

However, this is rendered null by the fact they are the sick fucks behind the Daemonculaba. Due to its nature, the women themselves may have the gene-seed organs implanted (among other things), or at least the DNA required to be able to quickly grow the organs into a child stuck in their wombs. While the poor, innocent, scared kid inside can become a skinless abomination/mutant, they will normally emerge a genetically stable, gene-seed uncorrupted skinless adult SPHESS MAHREEN with the mind of that same scared child. Suffice to say, this is a sick way of getting to FEMALE SPHESS MAHREENS, and is DOUBLE HERESY.

Luna Wolves/Sons of Horus (XVI Legion)

• Implicitly defective Ossmodula, causing increased cosmetic similarity to their Primarch.
• Implicitly defective Biscopea, causing increased cosmetic similarity to their Primarch.

No real defects, but the tendency for members to look more like their Primarch is more pronounced (and possibly more frequent) than it is for members of other legions to resemble their own respective Primarchs. "Little Horus" Aximand was the most notable example of this, but by no means the only one. This gives extra meaning to the "Sons of Horus" thing. In fact, before the Luna Wolves were renamed the Sons of Horus, Luna Wolves that resembled Horus in appearance were known as "Sons of Horus". Ten thousand years in the Eye of Terror has left the Legion's gene-seed more unstable. However, we do know that new Black Legionaries are inducted constantly (seeing as they are the primary generic Chaos Space Marines faction), so its a good bet a large part of theirs is still usable and that generations of children for millennia to come will be torn away from their parents and families, unfortunate enough to live in the Eye Of Terror, to be inducted into the Legion. Furthermore, given that they frequently "recruit" from other chapters and legions with the only real requirement being loyalty to Abbadon, it's not always easy to tell which Black Legionnaires are descended from Horus and which are from other Primarchs, outside of the aforementioned lookalike business.

Word Bearers (XVII Legion)

• Implicitly defective Catalepsian Node.

The Word Bearers suffer a unique "quirk" in their gene-seed, resulting in a psychologically obsessive loyalty to something, be it a man, a religion, or even an ideal. The Word Bearers in turn thus have a much stronger sense of loyalty to their battle-brothers and Primarch than other Chaos Space Marines, helping them to maintain cohesion and avoid treachery, but this loyalty also borders on fanaticism. Otherwise, while given to some instabilities, they have been able to maintain strength.

It has been suggested that Word Bearers have an extraordinary amount of control, both mentally and physically, far more so than normal Astartes. It's been show in their ability to directly control the processes of their internal organs and is attributed as the reason why Word Bearer Diabolists are fairing so well against the Neverborn.

Salamanders (XVIII Legion)

• Unique Melanchromic Organ, causing their skin to turn pitch black (so much so that it looks genuinely unnatural) instead of waiting for radiation to happen to fix it.
• Unique Occulobe, causing limited ability to see infrared emissions, referred to as "fire-sight". While not as potent as what an Auspex scanner would be capable of, it can be combined with specialized materials within armor and standards to produce decorations and tactical markings that only the Salamanders can see. This also gives them glowing red eyes, Which makes sense.
• Possibly altered Mucranoid - high degree of heat resistance.
• Possibly altered Biscopea or Lyman's Ear - tend to be somewhat slower moving than other marines, although sometimes it's blamed on the somewhat high gravity on Nocturne or the unusually methodical mindset of it's inhabitants, since they can still run at the same level as any other chapter. (Way back when, they had -1 Initiative.)

While slightly dysfunctional, their gene-seed is pretty stable. Would probably be, like the Iron Hands, a common source of new chapters if not for, again, their extreme decimation during the Drop Site Massacre. They've since recovered, perhaps a little quicker than the Iron Hands (possibly due to being a little closer to the center and being able to evacuate more marines, As well as still having their Primarch at the time), and while they claimed no official kinship with any successors before the Ultima Founding, there are at least a couple of examples with similar unit markings and tactical dogma.

Major Mutations

These Chapters feature at least one problem as least as bad as missing an entire organ, scaling up to the worst issue, where they go World Eaters-level crazy without needing any implants to get there (Blood Angels and Space Wolves).

White Scars (V Legion)

• Tendency to do whatever they fancy.
• Unrestrained love towards doing strange things to engines buffing their bikes and battleships to ride into battle while riding into battle.
• A defect called "Chogorian Savagery". Basically its diet Black Rage, born from when Mortarion almost killed their primarch during the Siege of Terra. This flaw appears in 2 stages:
  • Stage 1 - A Moment Unrestrained: The Battle-Brother's discipline begins to slip and falter, allowing brief moments of untamed ferocity to emerge in the heat of battle. Often, the Battle-Brother will not even know that he is succumbing to this savagery, and it requires the presence of his brothers to warn him of his failure.
  • Stage 2 - Suppressed Rage: Within the darkest reaches of the Battle-Brother's mind lurks the shadow of his rage and fury, seeking any opportunity to emerge, and causing the Battle-Brother to become belligerent and quick to anger.

Mantis Warriors

• Possibly Melanchromic Organ as all have green eyes and black hair of their Primarch.
• Malfunctioning Preomnor gland (possible explaining the cause of the gene-flaw of the other White Scars chapters)

Their mutation called "Battle Haze" causes their Preomnor gland to secrete a potent neuro-toxin that permanently changes the Astartes' physiology; the Space Marine is overcome by a feeling of the need for forgiveness for his many sins and shortcomings and a dramatically increased reverence for the Emperor; and increases his neurological reaction rate to that only achieved in near-precognitive states by most psykers and increases his already superhuman strength to a great extent. However, this physiological change is irreversible and the Astartes' sight becomes tunnelised to the point of no longer being able to notice anything in his field of view that is not a target or a foe. Much like the Death Company of the Blood Angels, the Mantis Warriors organize Battle-Brothers who have been lost to the Battle Haze into specialized units called "Mantis Religiosa".

Destroyers

• "Chogorian Savagery" turned up to 11. To the point even other White Scars question the stability of the Destroyer's geneseed. More evidence that the flaw stems for the gene-seed of the White Scars, and not from what population they are pulled from. Probably at least a bit psychological and genetic memory related, what with the most psychotic Scars forming this Chapter in the Second Founding.

Imperial Fists (VII Legion)

• Missing Betcher's Gland (no acid spit).
• Missing Sus-an Membrane (no suspended animation).
• Possibly increased pain tolerance and/or a tendency towards masochism.

Despite missing two organs, the Imperial Fists gene-seed is considered to be amongst the most stable and most manly sources. It is the second most desirable source of gene-seed for Foundings. Second to the Ultramarines, of course. The Fists however, display a constant need to punish themselves for perceived failures, both real and imagined. While this is barely noticable in the primogenitor chapter, several successors manifest more pronounced versions of this flaw. Missing Sus-an Membrane is also a big deal, as it removes a "safety net" most other chapters have, and most times when Imperial Fist "falls in battle" he actually dies for good instead of passing into coma from which he could be brought back later if marines win the engagement and recover his body. That's why Imperial Fists and all their successor chapters need multiple recruiting worlds. Silver lining though is that sons of Dorn don't run the risk of being captured by Chaos and brainwashed if they fall in battle and their body is then recovered by forces of Chaos instead of allies.

• • Complicated by the War of the Beast, in which the Imperial Fists were literally wiped out to the last man. In the aftermath, their successors got together to donate gene-seed to reconstitute the chapter, making their post-Beast gene-seed a chimeric melting pot of separate mutations and oddities. Additionally, it's not confirmed whether the mechanicus were holding and offered stocks of original, untainted gene-seed as part of this process. Even Primaris marines who were made with the original stuff (and in some cases are original legionaries) may be different as it's not clear what effect Primarisification has on the gene-seed or if it's possible to make firstborn from it. In short; the Imperial Fists gene-seed may be unique among loyalist chapters in that it simply no longer actually exists.
    • Except, hilariously, in some Iron Warriors, who take a particular spiteful delight in stealing their arch-nemeses' progenoids.

The Excoriators

• A mutation with some similarities to the Blood Angel's Black Rage, known as "Dorn's Darkness.". The affected appear, to the untrained eye, as merely a wretched palsy: a slackness of the jaw, a tremor of the limb, a blankness of the eye. However, reports from the affected say the experience is a living nightmare, a sleeping wakefulness in which they relive the bottomless woe and paralyzing fear of Dorn's darkest moment: the moment he saw the Emperor near death after the battle aboard the Vengeful Spirit. When affected by Dorn's Darkness, its victims cannot speak or communicate. They cannot feed themselves or take water and seem feverishly insensible to everything happening about them. Basically, if the Black Rage is unbridled RAGE unbound, Dorn's Darkness is crippling, ruinous depression.

Crimson Fists

• A mutation similar to Dorn's Darkness, known as "Only Honour In Death". The "curse" causes a massively increased willingness to fight to the last, holding their ground with a siege-like mentality which will see them triumph or die in the attempt the more foes that the Astartes faces, and the longer they are in combat, the more likely and stronger the curse becomes, making them feel emotions much like what Dorn felt during the Iron Cage Incident. When succumbing to the "curse" the affected Battle-Brother sees only death: the death of their comrades, the death of the world, and the death of their foes. The curse is seen to come in three stages:
  • Stage 1 - No Retreat. As the odds mount so does the Battle-Brother's stubborn refusal to retreat, seeing only the glory and honour in defeating such overwhelming foes if even the barest glimmer of victory remains. The Battle-Brother will even continue to fight, even if the Codex Astartes does not support that action, until there is no hope of victory left.
  • Stage 2 - Suffer not Defeat. The Battle-Brother would rather die fighting than accept that he has failed. This can lead to reckless behaviour as he throws himself into the fray or takes on excessive odds to win when prudence and tactical reason would dictate he withdraw. Basically a mini-Black Rage
  • Stage 3 - Unto Death. The Battle-Brother gains a fatalistic desire to sell his life for the cause and will choose to expend his own life for a chance at victory or the safety of his brothers. He has come to accept that perhaps only the cost of his own life will be enough to secure victory. This can have dire consequences should the circumstances present him with a chance to martyr himself for the Emperor and bring ruin to the enemies of the Imperium. This means he will fight on regardless of his wounds and continue to attack or place himself in the path of attacks even though the next blow could mean his end. He will also gladly accept suicidal or near-suicidal tasks.

Space Wolves (VI Legion)

They actually have two different implanted DNA sources: the second is the Canis Helix, which makes them viking werewolves (long canines, tough skin, awesome sense of smell, animalistic RAEG in combat) or, if one is really unfortunate, into a giant wolf (the ones they ride at that), and their super-unstable geneseed, which can't function without pre-implanted Canis Helix and has been found to be incompatible with any populace outside of Fenris. For the most part, they get around this by just founding tons and tons more Space Wolves than the Codex Astartes would normally permit, meaning that they still tend to have tons of chapters' worth of marines on call. Of particular note are the Wulfen, who are Space Wolves who happened to be in the Warp for over ten thousand years. Instead of being mutated into unmentionable gribbly beasts, they managed to turn into literal werewolves with savage strength, meaning there's a possibility the gene-seed has a factor that lets them resist the influence of Chaos.

While many factions within the chapter are proud of how "potent" their gene-seed is, plenty of Space Wolves have constantly labored to stabilize their genome in order to found successors to cordon off the Eye of Terror, and crusade against the Thousand Sons, a plan called "the Sons of Russ." Once, they got really close, which Magnus the Red didn't like, so he took his Marines and bumped off the Wolf Priest in charge (and generally wrecked things for the Space Wolves), using some Tzeentchian trickery to get an opening. In short, he got shit done!.

Wolf Brothers

See Space Wolves above turned up to 11. As in the entire chapter devolved into werewolves.

Night Lords (VIII Legion)

• Defective Occulobe (causing them to have black irises & sclera, enhanced night vision but more sensitive to bright light.)
• Malfunctioning Melanchromic Organ (their skin is always pale and their eyes are always dark. Much like the Raven Guards.)
• Occasional visions of the grimdark future, even if not a psyker

The Night Lords are officially the most stable of the Traitor Legions, a trait often put down to the fact that they rarely dwell in the Eye of Terror. That, and they don't even like Chaos or the gods. What little mutations there are are usually upon their Sorcerers...what ones have survived do not have them anymore either. There is one defect on Curze's Geneseed, and that is the Occulobe defect, causing them to have black irises & sclera, and astonishing sight in the dark, even more than a normal space marine. Although, there is a bad side of this effect: Light hurts their eyes like hell, and blinds them for a short time. Also, Curze's gene-seed may have passed onto psykers within his Legion his "gift" of seeing the future in the worst way possible, and as scenarios that become self-fulfilling prophecies. In one case, it was found out that the Primarch's "gift" was actually the side effect of a marine's body slowly rejecting the gene-seed. Prior to receiving this gift, he never demonstrated any psychic capability. Even so, this situation is rare.

Blood Angels (IX Legion)

• Malfunctioning Omophagea implant (Occasional bouts of temporary bloodthirsty, genocidal rage, otherwise known as the Red Thirst.)
  • Occasional bouts of permanent, bloodthirsty, hallucinatory, genocidal rage, better known as the Black Rage.
• Extreme longevity.
• Can turn even mutants to Astartes, extremely adaptable.

Stable, but flawed. Prior to the Horus Heresy, Sanguinius discovered that some of his sons were beginning to suffer the Red Thirst, where they would descend into a thirst for blood, whether human or xenos, in a blind rage. If that wasn't bad enough, as a result of Sanguinius's death at the hands of Horus imprinting itself on their genetic memory (or something), all Blood Angels and their successors have the Black Rage; a berserk rage where they believe themselves to be Sanguinius during his final battle. Meaning that Fucking Horus quite literally hit Sanguinius so hard that his grandchildren's-grandchildren's-ad-infinitum-children still feel it. Notable due to the age of the flaw it means that it does not register on genetic purity test since, as a first founding legion, any additional geneseed would be compared to an already 'tainted' baseline.

Angels Encarmine

• See the Blood Angels above, but with the Black Rage turned up to 10

Flesh Tearers

• See the Blood Angels above, but with the Black Rage turned up to 11.
• Also Red Thirst turned up to 10.

Knights of Blood

• See the Blood Angels above, but with the Black Rage and Red Thirst turned up to 12 (and the dial snapped off).
  • It got so bad that they were declared renegade and not refounded after they sacrificed themselves to save the Flesh Tearers at Baal.

Blood Drinkers

• Malfunctioning Mucranoid (skin glands atrophy, causing the pores to close. Named the "Weaver" by the Blood Drinkers)
• See the Blood Angels above, but with the Red Thirst turned up to 10. Due to further mutation of the Omophagae implant.
• Black Rage turned down a notch, but this isn't genetic. The entire Chaplaincy is in on a Daemonic pact to reduce its impact.

Lamenters

• All the problems of Blood Angels (tho it took them longer to show).
• Being overly emotional sad sacks, but this is as justified as it gets because...
• Unlucky as all fucking hell. Best way to describe it is fate itself is just constantly giving them the middle finger. This might have something to do with them coming out of the Cursed 21st Founding.

Thousand Sons (XV Legion)

• Increased chance of bringing out psychic powers, as well as amplified psychic power.
• Recipients subject to a high rate of mutation, called The Flesh Change; unstable.

The most unstable genome in the Space Marine legions, the Thousand Sons suffer from the Flesh-Change, a problem that caused Thousand Sons to mutate spontaneously. Magnus the Red fixed it, with some "help" from Tzeentch. After the Horus Heresy, the flesh-change came back with a vengeance. Ahzek Ahriman tried to fix it with his Rubric, and he did end up stopping the flesh-change... by turning all non-psyker Thousand Sons into dust; sorcerers also had their powers enhanced in addition to no longer suffering ill-effects from the Flesh-Change. So while theoretically, the Sorcerers of the long suffering Legion can provide gene-seed, in practice, it's corrupted so often that few will ever make new Marines. Thousand Sons specifically hunt for healthy and stable psyker boys to implant them with their oh-so-rare gene-seed to ensure the recipient would become a sorcerer. If "Masters, bidding!" TS character words to be trusted, Thousands Sons found a way to implant their progenoids into grown up men (so they can teach and test them for longer before implantation), and perform a scaled down Rubric on them to prevent the Flesh Change. According to Phil Kelly they also replenish their numbers by summoning back souls of destroyed Rubrics, and Magnus supposedly can bring back Sorcerers he isn't pissed off at, so gene-seed wouldn't matter much to them. It is known that Magnus the Red used his knowledge of Biomancy to turn his mentor Amon into an Astartes despite his old age, so its possible this technique was passed down to his Legion.

Raven Guard (XIX Legion)

• Malfunctioning Melanchromic Organ (their skin is always pale and their eyes are always dark. Anti-Salamanders, basically.)
• Missing Mucranoid (no super-sweat).
• Missing Betcher's Gland (no acid spit).
• Occasional bouts of suicidal rage (Luckily for them this is not permanent unlike a certain other chapter.)

While their primary gene-seed is decently stable, when under extreme stress a percentage Raven Guard warriors will lapse into bouts of suicidal rage known as Ash Blindness or Sable Brand. This is rarely permanent, but occurred with enough frequency that Corax would organize these warriors into the Moritat. Their missing and malfunctioning geneseed organs is a result of Corax, out of desperation following the Drop Site Massacre, delving into use of forbidden lore and archeo-tech from days long past to accelerate gene-seed maturation, and try to boost his Legion's numbers after their losses during the Horus Heresy. He was successful, until the Alpha Legion decided to make the whole project go FUBAR. The subsequent 'Marines', if they can be called that, were numerous, but were barely controllable monsters and/or would simply die off as a result of the extreme imbalances in their dispositions. Presumably, a few of them would even have functioning Progenoids, let alone usable gene-seed. Due to the law of averages though, at least a FEW would have had to been stable enough, and is being worked on secretly by the Raven Guard Apothecaries and associated Mechanicus Biologicans. Otherwise, despite being pretty fucked up by the Drop Site Massacre, they've mostly recovered (thanks in no small part to having the only loyalist Primarch to get away from the Massacre in one piece) and do still contribute to successors here and there.

Raptors

• Same as the Raven Guard, except they have functional Bletchers Glands (making them more like another black-eyed / pale skinned gene-line).

Death Spectres

• Similar Melanchromic Organ as Raven Guard, though a specific shade of albino and their eyes universally glowing red (basically albino Salamanders).
  • also gives them a 100% certain chance of alopecia. (They are genetically bald From the day they were born till the day they die)

Marines Errant (Ultramarines Successors)

• Suffer a psychological flaw called "Crusaders Call", which makes them unable to refuse the call for a crusade. This flaw manifests in three stages:
  • Stage 1 - Unending Valour: The Battle-Brother has become filled with pride in the history of his Chapter's bravery and devotion and sees his own existence as a vital part of continuing its traditions. When his squad is required to do its duty he will make sure that it is carried out, squashing any doubt his fellow Battle-Brothers might have or any weakness they might show in the face of honour.
  • Stage 2 - Duty through Sacrifice: True duty can only be proven in death or an equally potent sacrifice, a fact which is known to all true servants of the God-Emperor. The Battle-Brother knows that to prove himself he must be the one to stand in the way of the blade or shield his comrades from the blast, so that there can be no doubt about his devotion or his loyalty to the Emperor.
  • Stage 3 - Lead the Charge: In boarding actions, the greatest honour goes to those first through the breech and first to face the foe. The Battle-Brother accepts that he must take this role as his honour demands it, and it is just another way to prove his courage and loyalty in the eyes of his peers. If there is a choice or a chance to lead a charge or be the first to face the enemy, the Battle-Brother must take it. Furthermore he will seek out such opportunities, volunteering for dangerous duties if they mean he will get to lead others into the fray. In combat, the Battle-Brother will also seek out the most dangerous or powerful-looking foes, even at the cost of ignoring closer or more pressing enemies to his brethren's tactical disadvantage.

It is unclear if these are actually geneseed flaws or cultural issues ascribed to geneseed

Soul Drinkers

• Enhanced Omophagea organ (In addition to gaining the consumed creature's memories, the Soul Drinkers could also experience the creature's emotions during the events recorded in their memories as a result of their mutation). This is despite being a 2nd Founding chapter, such that their gene-seed shouldn't have had flaws like that. And the gene-seed was enhanced, not degraded....

Blood Ravens

• A mutated Catalepsean Node - gives them perfect memory recall, but unable to enter REM (deep) sleep.
• Increased chance of bringing out and enhancing psychic powers.
• Small tendency towards mutations (hurr sounds like any other Legions we know of?).
• Horrendously massive tendency to develop kleptomania.
• The pores in their scalps secrete a substance not all too dissimilar to hair gel, often resulting in a higher tendency to develop hair-etical hair compared to other chapters.
  • Alternatively, the pores on their scalps may prevent hair from growing altogether, giving the marine Astartes-pattern baldness. It has been long thought to be a terrible weapon to behold.
• Unknown mutation referred to as "unavailable voice actor syndrome" that results in their voices changing every few years.

Carcharadons

• Malfunctioning Melanchromic Organ (pale skin and black eyes like the Night Lords and Raven Guard)
• Mental deterioration called "Chill of the Void". This deterioration proceeds like a mini-version of what Konrad Curze went through as he became the Night Haunter. This deterioration occurs in three stages:
  • Stage 1 - Coldly Formal: The Battle-Brother withdraws even more from the company of his colleagues, taking refuge in a chill and formal demeanour. In addition, the Battle-Brother tends to speak almost entirely in High Gothic when addressing a non-Carcharodon, although they can speak in Low Gothic if need be (but they really do not want to).
  • Stage 2 - Merciless: The Battle-Brother does not tolerate any enemy, whether xenos, Heretic, or even fellow humans who may have been misled into rebellion against the Imperium's rightful rule. The Battle-Brother grimly executes any opponent he encounters, even if they have surrendered, possess valuable intelligence, or are not front-line combatants.
  • Stage 3 - Silent as the Depths: The Battle-Brother withdraws almost entirely from interacting with others. He will not lead squads and will almost always carry out missions on his own.

Black Dragons

• Extreme Ossmodula defect. causes them to grow bone spikes all over. Including full on Wolverine type claws from their arms.
• Moderate Betcher’s Gland defect. Their mouths, both inside and lips, are usually black in colouration. So too, their tongues, which taper to a sharp point like a dagger.

Also causes purple venom to be secreted from their teeth (which resemble fangs) and their tongue.

Flame Falcons

• Spontaneous combustion into warp fire.
• Increased abilities from said spontaneous combustion instead of horribly burning to death.

Minotaurs (21st Cursed Founding-type)

• ABSOLUTELY BATSHIT GOD DAMN ASSFUCK INSANE

These guys had....problems....big problems. They'd basically wipe their asses with any battle plans made by other Imperial forces, before bee lining it to the nearest horde of enemies (not bothering with any of that pussy shit like "recon", "intel", or "even bothering to look out the window at the enemy") and rushing into melee ranges to RIP AND TEAR. But, this doesn't apply to the original and current units. You see, the Minotaurs chapter have a good chance of actually being a cover to get a loyalist (they had missed out on the Heresy entirely due to their distant posting and spent the interim years unaware of the situation holding the frontier) Iron Warrior garrison unit rediscovered in the Great Scouring legitimized by the High Lords of Terra. See their article (Minotaurs) or the Traitor Legion Loyalists article for further details.

Sons of Antaeus

• Unknown mutation that makes them exceptionally durable and tough. To the point of easily being a match for the Death Guard (who are boosted by Chaos energies).

Storm Giants

• Unusually physically strong, even by Astartes standards. Speculated to be from a mutation or alteration in their Biscopea.

A Note On That Sick Fuck Fabius Bile

Fabius Bile is fucking insane. I mean shit, his labcoat is made of human ASTARTES skin - normally only Space Wolves and Night Lords wear clothing made of that stuff. While originally the Chief Apothecary of the Emperor's Children, he is now a freelancer. His goal even before the Heresy was to find and unlock the secret to the creation of the Space Marines. In the Warp, he provides a valuable service to the Chaos Legion warbands and their auxiliaries of billions more regular Chaos humans - while often millions strong even now, the warbands still need to replenish their numbers. Considering how many die and how often they are away from any place where they can produce more, that's a lot. Even with slave raiding into the Imperium, child abduction both in and out, breeding centers (stop touching yourself), and many other creative ideas, it's still not enough. Fabius provides services of human cloning, along with gene-seed production, among his many skills. As he is not a super-devotee of Chaos, the gene-seed he has and makes is some of the least corrupted found within the Eye Of Terror. Of course, he fucks this all up by hopping up his own warband's clones with the same psycho drugs and shit that make monsters out of men. Lore hints that his genetic tinkering is far more than we have been lead to believe - much of his 'changes' to even the standard human genome are genetic and now passable by both men and women. Even now, the Imperium is trying to hunt and eradicate what they call his 'New Men'. He also has access to a small quantity of what is alleged to be the blood of the Primarch Sanguinius. Why did he get that last part? To clone the Emperor. Never let it be said that this fuck, sick as he may be, lacks ambition.

21st Founding

Over the millennia, further mutations have afflicted the many Successor Chapters, but the 21st or "Cursed" Founding deserves a special mention, because a bunch of tech-priests decided that they wanted to try and fix the flaws, but failed so hard they actually created some new ones.

• Bursting into Flames: Yes, the Flame Falcons really were covered in fire. The Flame Falcons thought it was a blessing from the God-Emperor of Mankind, but the Inquisition disagreed, and sent the Grey Knights to wipe them out.
• Bone Growths: Due to a malfunctioning Ossmudula, the Black Dragons suffer a bone solid growth to harden and extend from their forehead and forearms during times of stress. The ones on the forearm can be used as weapons against unarmored opponents, so they decided to coat them in adamantium. So in essence, they're space-Wolverine-after-the-bone-claws kind of guys. Issue is that this is a very visible mutation, and as this is the grimderp Imperium, mutations from the sacred human form are EVUL, even if they are fucking awesome.
• Indestructibility: The Sons of Antaeus are much tougher than normal marines, and nobody knows why. Some speculate something about a super-reinforced skeleton, some say it's Nurgle's doing, etc. The only real truth to it is that there's some sort of mutation present in the gene-seed.
• Bad Luck: The Lamenters chapter's modifications seemed to have fixed the Red Thirst and Black Rage, but as a trade off they seem to be cursed with some of the worst possible luck, essentially giving up the Khornate Curse for a Tzeentchian one. Many other chapters of the 21st Founding were also cursed with bad luck but the Lamenters are the only known ones that didn't get wiped out by it so that may say something about how badass they are.

A Note On Implant Organs/DNA Elsewhere

While the SPHESS MAHREENS are the most widespread users of artificial glands, organs, and such, they are not the only ones. Inquisitors likely have access to organs or other artificial mutations they require thanks to their influence in the Imperium. Depending on what world they hail from and its beliefs, Imperial Guard regiments and PDF may also have artificial organs, or even whole new ones derived from millennia of mutations, which of course has certain advantages. Furthermore, Warp Exposure can result in a change of the functions of organs - a simple example of this is the Cadians, who nearly universally have powerful, violet colored eyes. Gland Warriors were an Adeptus Mechanicus project to help legions of Guardsmen heading into battle against a Tyranid hive fleet. They were to do battle in an incredibly toxic and hard environment, and a variety of experimental organs and DNA strands were utilized - including organs that would secrete stims, painkillers, and various medical drugs, more powerful natural air filters, and the like. While only three survived the whole campaign in the end, the AdMech was pleased and took the three to be debriefed.

It is also noted that Gene-seed can be decoded, but it requires eons of knowledge about biology, medicine, and so on. Only the Emperor or AdMech, with their advanced equipment, is able to do so (and even the AdMech couldn't fully work it out). The hard part is to decode the "impossibly complex" amino acid chains of the gene-seed (which FYI makes no sense, since DNA is encoded in nucleic acid base pairs, and amino acids are used to construct proteins but whatever) and extract the necessary information encoded within the Gland's stored zygotes. Arik Taranis was able to decode it in a dusty, old, half-broken lab—with a box of scraps—using the knowledge gleaned by just watching the Emperor work, and remade it to fit in his Thunder Warrior body.