Archive for effective population size

You Paid for 415 Kilos of Genetic Gain. Inbreeding Kept 143 of Them.

Italian researchers tracked 27,735 Holsteins for five years. The breed earned 415 kilos of genetic gain per lactation. Inbreeding took 143 back. Do you know your own herd’s number?

Executive Summary: Ablondi’s Parma team measured the same 27,735 Holsteins twice, and pedigree said 7% inbred while the genome said 16% — with the milk loss tracking the bigger number, 61 kg per point against 44. That gap matters to anyone genomic-testing females and still running matings off pedigree EFI, because you’re managing to the instrument that reads low. Across 2015 to 2020 the Italian Holstein banked 415 kg of genetic gain per lactation and inbreeding depression took 143 of it back, though the authors are careful to say the breed still kept 272 kg net; the worry is the trajectory, not a bill already due. Run Pryce’s scaling on your own operation — 0.4 to 0.6% of your rolling herd average per point of inbreeding — and a decade at the U.S. rate of 0.36 points a year costs a 12,000 kg herd 170 to 260 kg per cow, per lactation. North American Holstein’s genomic-era effective population size sits somewhere between 24 and 42, and even the optimistic end is under the FAO’s critical threshold of 50. Four major studs built internal elite-female programs between 2008 and 2015, only Select Sires has published a sourcing split, and none of the eight organizations we contacted responded — so nobody outside those companies can say where your next bull came from. The tools that fixed this in the 2000s are still available; the open question is whether anyone uses them at a 2.2-year generation interval, and whether you’ve ever seen your own two numbers side by side.

Michela Ablondi’s team at the University of Parma did something most inbreeding studies skip. They measured the same 27,735 Holstein cows twice.

Once on pedigree — the way most mating programs still calculate it, working back through recorded ancestors. Once on genome, marker by marker across 84,443 SNPs. Pedigree came back at 7% average inbreeding. The genome came back at 16%.

Then they checked which number the cows agreed with. Each 1% rise in pedigree inbreeding cost 44 kg (97 lbs) of 305-day milk. Each 1% rise in genomic inbreeding cost 61 kg — 134 lbs. Same animals, same lactations, and the depression tracked the bigger figure.

That’s the part worth sitting with, because Lactanet’s August 2025 figures put average inbreeding for Canadian Holstein heifers born in 2024 at 9.99%, climbing about a quarter point a year on a pedigree basis. If pedigree is the instrument reading low, then the number most producers are watching is not the number doing the damage.

Two Instruments, Same Cows, One Reads Low

The correlation between the two measures came in at 0.65. Related, nowhere near interchangeable. The reason matters: pedigree can’t see Mendelian sampling, and it’s only as good as the records behind it. Ablondi’s group had 24 generations to work with and a complete generation equivalent of 10.6 — deep by any standard — and pedigree still read less than half what the genome showed.

Your own two numbers won’t be 7 and 16 exactly. Ablondi’s herds had unusually complete records, and the size of your gap depends on your pedigree depth and how much recent breeding runs back through the same bulls. What won’t change is which direction the error runs. Pedigree reads low, every time.

Ablondi’s group also split inbreeding by age. Runs of homozygosity longer than 8 Mb — the fingerprint of recent inbreeding — carried the negative effect across every trait. Pryce and colleagues found the same thing in Australia a decade earlier with 8,853 Holsteins and 4,138 Jerseys: ROH over 3 Mb cut milk yield and captured recent inbreeding independently of overall homozygosity. Two teams, two continents, ten years apart. Recent inbreeding is what hurts.

In the U.S., Steyn and colleagues documented the annual increase accelerating from 0.11 percentage points a year between 2000 and 2008 to 0.36 between 2013 and 2021, working from CDCB data in the Journal of Dairy Science in 2022. Three continents, same direction.

The 143 Kilos You Paid For and Didn’t Keep

Now the cost at breed scale. Between 2015 and 2020, the Italian Holstein’s pedigree inbreeding climbed 2.35%. Realized genetic progress for milk came in at +415 kg (915 lbs) per lactation.

Inbreeding depression took back 143 kg — 315 lbs. That’s 34.5% of the gain.

Be precise about what that means, because the authors are. Their own conclusion is that “the realized genetic progress has so far successfully counterbalanced the negative effect of inbreeding” for milk yield. The breed kept 272 kg net. Progress didn’t stop, and anyone telling you the Holstein is going backwards isn’t reading the paper.

What worries Ablondi’s group is the trajectory. The tax compounds, it lands on fitness traits this study didn’t even measure, and inbreeding is projected to climb faster in the generations ahead. Run the same arithmetic on the pedigree measure and you get a smaller bill — 103 kg, 24.9%. Two instruments, two answers again, and only one of them is telling you what your matings actually cost.

Compare the extremes and it gets uglier. Moving from the 5th-lowest to the 95th-highest inbreeding percentile cost 263 kg (580 lbs) per lactation on pedigree — 2.4% of the herd average — and 561 kg, 1,237 lbs, on genomics, or 5.2% of the herd’s 10,836 kg (23,890 lbs) average. Fat dropped 1.31 kg and protein 2.45 kg on the genomic measure.

What nobody can measure is which bulls are driving it, because no public database records where an AI sire came from.

Below the Line Where Fitness Erodes

The FAO puts the critical effective population size at 50. Not 50 cows in a barn — 50 in the genetic sense, the number of breeding animals that would produce the inbreeding rate you’re actually seeing.

Makanjuola and colleagues measured it for North American Holstein and Jersey using Canadian Dairy Network records on 676,594 Holsteins, publishing in the Journal of Dairy Science in 2020. Across 1990 to 2018, Holstein Ne came in at 66 on pedigree and 43 to 46 on genomic measures. Jersey held up better at 64 to 85.

Narrow to the genomic-selection era and the rate of inbreeding per generation climbs to between 1.19% and 2.06%, depending on the measure. That puts Holstein Ne somewhere between 24 and 42.

Which is why you’ll never see an honest single number here. Pedigree and genomic measures disagree on the rate of inbreeding per generation by nearly a factor of two, so North American Holstein Ne isn’t a dot — it’s a corridor running from 24 to 42, and both ends sit under the FAO line. Same argument as the 7-versus-16 gap, one level up.

Behind it sits a generation-interval collapse. Holstein sires of bulls averaged 8.08 years in 1990 and 2.32 by 2016 — a 71% reduction. Guinan and colleagues documented the same trajectory in U.S. data on the sire-of-bulls pathway, 7.1 years down to 2.2.

Nordic populations get held up as the counterexample, and the picture is genuinely mixed. Tenhunen and colleagues found Nordic Holstein Ne falling from 54.3 before genomic selection to 42.8 after, with future Ne dropping from 198.8 to 42.7 and female coancestry rising from 0.02% to 0.39%. Their conclusion: Nordic Holstein “faces more pressing concerns.”

But read that comparison carefully. At 42.8, Nordic Holstein sits at the optimistic end of North America’s 24-to-42 corridor — better than the pessimistic read, roughly level with the generous one. Methods and study periods differ between the two papers, so treat it as directional, not a scoreboard. Nordic Jersey held essentially flat at 29.4 before and 30.3 after.

Against the FAO line the pattern is blunt. North American Holstein on genomic measures, Nordic Holstein, and Nordic Jersey all sit below 50. North American Jersey is the only measured population that clears it.

Can Anyone Tell You Where Your Next Bull Came From?

Not from anything published.

Every major stud runs an internal elite-female program now, and most say so openly. Select Sires launched ART in 2008, targeting Holsteins “genetically unique compared to those found in the general population.” Alta started PEAK in 2012 as its genetics development arm during a restructuring built around genomics, and PEAK joined URUS when that holding company formed in October 2018 — PEAK’s own materials date the entity to 2018. Semex founded Progenesis in 2013, calling it “Semex’s industry-leading internal dairy breeding program.” ABS Global shipped its first commercial units from an elite female nucleus herd in 2015.

Look at the dates. 2008, 2012, 2013, 2015 — every major stud built an internal elite-female program inside a seven-year window, and that window opens the year before USDA-AIPL published the first official U.S. genomic evaluations in January 2009. Four companies didn’t independently arrive at the same structure by coincidence. They read the same new tool the same way.

Four programs. One published split.

Select Sires puts roughly 40% of the Holstein bulls it samples each year as ART graduates, with 60% acquired from producers across the U.S. and Canada. That figure appeared in a January 2024 company post, and ART program manager Mark Kerndt repeated it on the record in a July 2025 Select Sires podcast, describing an internal female donor program running since 2008 and transferring roughly 15,000 IVF embryos a year. Two independent company sources, one of them a named executive speaking on tape.

Credit where it’s owed. That’s the only proportional sourcing figure any of the four has put in public, nobody made them do it, and it’s now better documented than anything else in this section. ABS, Semex and PEAK describe their programs without quantifying what share of released sires come through them — which leaves the one company that disclosed as the only one whose disclosure can be picked apart.

Semex says the Progenesis team “continues to add bloodlines that diversify the program.” That sentence is the whole problem in miniature. It’s substantive, it’s checkable in principle, and it’s unverifiable from outside. Semex may be entirely right. Nobody outside Semex can confirm it.

Past what companies volunteer, the trail goes cold. NAAB’s Semen Sales Report tracks units by breed and category, and we found no field recording origin. CDCB publishes proof runs and reliability figures on every sampled bull in the U.S. system, and we located no public output sorting sires by nucleus, contracted, or open-market origin.

The Counterweight Nobody Expected

Here’s what complicates everything above.

United Sires — NAAB codes 596 and 796, registered with NAAB in February 2024 — is an independent breeder partnership with no ownership tie to any of the majors. By The Bullvine’s count of Holstein Association USA’s top 200 TPI genomic young bulls — 85% genomic reliability minimum, from the CDCB run published April 7, 2026 — it holds 30 of them, 15%. That’s third behind Select Sires’ 59 (29.5%) and Semex’s 40 (20%), and ahead of ABS at 12, Alta at 11, GENEX at 9 and CRV at 8.

We’re citing April rather than the August 2026 run on purpose. April is when Holstein Association USA changed the TPI formula, shifting production weights from 19% protein and 19% fat to 24% protein and 14% fat, and it’s the snapshot that isolates what that change did to the standings. Bulls moved as much as 125 points in either direction without a single new daughter. Same bulls, same daughters, different instrument — the third time in this article that swapping the measuring tool has swapped the answer. Holstein Association USA has since published an August 2026 list, and the shares will have shifted some with it; treat the April figures as the post-formula-change baseline rather than today’s standings.

So whatever’s happening to the independent breeder, “shut out” doesn’t cover it. The pipeline is concentrating and a two-year-old breeder-owned partnership just took 15% of the top of the list. Both are true. Neither reconciles without the origin data nobody publishes — if the nucleus programs are as dominant as their scale implies, United Sires shouldn’t rank where it does.

Worth knowing what the modelling said before any of this got built. Dekkers and Shook, at Wisconsin-Madison, modelled nucleus schemes for commercial AI firms in the Journal of Dairy Science in 1990 (73:1920). Their finding, replicated since and echoed in Roden’s 1994 review: opening a nucleus raises both the intensity and the accuracy of female selection, and open systems deliver a higher rate of genetic gain than closed ones. They also found closed schemes could stay competitive at large nucleus sizes — so the easy version, closed bad and open good, is only half right. Design matters more than the label, which is exactly why the design should be public.

What It Costs, Country by Country

Four populations have been measured for inbreeding depression. Every one is negative:

StudyPopulationPer 1% inbreeding increase
Ablondi et al. 2023Italian Holstein, 27,735 cows−61 kg 305-day milk (ROH); −44 kg (pedigree)
Doekes et al. 2019Dutch Holstein-Friesian, 38,792 first-parity cows−36.25 kg (ROH); −37.95 kg (pedigree); −48.07 kg (genomic relationship)
Bjelland et al. 2013U.S. Holstein, University of Wisconsin−20 kg 205-day milk (ROH); −47 kg (genomic relationship); −53 kg (percent homozygosity)
Pryce et al. 2014Australian Holstein and Jersey, 12,991 genotyped cows−0.4% to −0.6% of phenotypic mean for milk, fat and protein

Figures as published. Multiply kg by 2.205 for pounds — 61 kg is 134 lbs, 36.25 kg is 80 lbs, 20 kg is 44 lbs.

Pryce’s framing is the one to run on your own operation, because it scales to whatever you produce. At 0.4 to 0.6% of the mean per 1% of inbreeding, a decade at the U.S. rate — 0.36 points a year, 3.6 points total — costs 1.4% to 2.2% of your herd average. On a 12,000 kg (26,455 lbs) herd, that’s 170 to 260 kg — 375 to 573 lbs — per cow, per lactation.

Ablondi’s team costed it in terms their readers feel immediately. Eighty percent of Italian milk goes into cheese, so they priced the loss in Parmigiano Reggiano — the country’s second-largest PDO consortium, 160,097 tonnes in 2022. Between the most and least inbred 5% of cows, the difference came to over half a wheel per cow per lactation on pedigree and more than a full wheel on genomics. Roughly €310 and €600 — about $360 and $700 USD, or $495 and $960 CAD, at 24-month-aged prices.

Pryce’s group mapped the damage to specific regions, finding stretches that cost up to 260 L of milk and 12.5 days of calving interval when fully homozygous. Ablondi’s team saw the same chromosome by chromosome — 16 of 29 autosomes carried a measurable hit to milk yield, with BTA14 the worst at −3.53 kg per 1% of homozygosity on that chromosome alone.

Bjelland’s U.S. work adds a cost that usually gets filed under calving management: stillbirths rose 0.25% for bull calves and 0.20% for heifer calves per 1% increase in inbreeding.

Doekes explains why recent matings matter most. New inbreeding — alleles identical-by-descent for the first time — cost 2.42 kg (5.3 lbs) of fat per 1%. Ancestral inbreeding cost essentially nothing, +0.03 kg. The damage lives in the matings you’re making right now.

The Tools You’re Probably Underusing

ToolWhat It MeasuresData Basis
CDCB Expected Future Inbreeding (EFI)Population-level inbreeding impactPedigree relationship to heifers born in the past 48 months; adjusts PTA accordingly
Genomic Future Inbreeding (GFI)True genomic relationshipDirect DNA marker overlap; consistently higher and more realistic than EFI
Lactanet Inbreeding CalculatorMating-specific inbreeding and parent averagePedigree-based; flags haplotypes and genetic conditions (LactanetGen.ca)

The EFI-versus-GFI choice isn’t academic. Makanjuola’s team found the correlation between pedigree and genomic inbreeding in Holsteins weakening materially across the period they studied — pedigree is getting worse at tracking reality, not better. Pryce’s conclusion was blunt: genomic measures can replace pedigree for estimating depression, using either the genomic relationship matrix diagonal or the proportion of homozygous SNPs.

Genomic-testing your females and then mating off pedigree inbreeding means managing to the number that understates the problem. That’s the whole finding, in one sentence.

One stud has published constraint evidence. C. Sun and colleagues at STgenetics presented Chromosomal Mating at ADSA in 2019 — a linear-programming system built on CDCB genomic relationship files covering 5,731 bulls and more than 1.3 million females, against a pedigree database of over 5 million animals. They reported average progeny predicted producing value up 17% and inbreeding down 30% against random mating.

Their design choice is the interesting part. Rather than setting a hard inbreeding ceiling, the system prices inbreeding as an economic penalty inside the optimization — a defensible way to handle a variable nobody has set a herd-level limit for. It’s also a company-authored conference abstract, seven years old and not independently validated, which is how you should read it.

Two Ways to Buy Pedigree Distance

One route stays inside the breed, and there’s a working example of it in Quebec. Blondin Sires — NAAB code 799, owned by Simon Lalande, Dann Brady and Nicolas Lalande — grew its share of Canadian sire usage from 2.8% in 2022 to 4.9% in 2023, a 75% jump in one year per Lactanet market data reported by The Bullvine in February 2026. The pitch is separation from the mainstream stack while staying Holstein; they prioritize proven performance in both conformation and productio, and use different sires than those targeting the top TPI or NM$ lists.

Be sure to check out our article with Dann Brady and team on their breeding strategy tomorrow. 

United Sires’ 15% share of the top 200 says there’s more room in that lane than the concentration story implies. The trade-off is real, though: independent-sourced bulls may carry lower reliability figures, and you’re still drawing from a gene pool whose effective size sits under the FAO line.

Going across breeds has a decade of peer-reviewed production data behind it. VikingGenetics’ ProCROSS rotates VikingHolstein, VikingRed and Coopex Montbéliarde, and a ten-year University of Minnesota study led by Les Hansen with Amy Hazel and Brad Heins measured it in high-performance Minnesota herds. ProCROSS cows returned 33% more lifetime profit and more daily profit — 13% for two-breed crosses, 9% for three-breed. Forty-one percent reached a fourth lactation against 21% of the Holsteins.

They also stayed 147 days longer, though Hansen’s own conference presentation of the combined two- and three-breed groups puts that at 153. Conception rate to first service in second and third lactation ran 45% against the Holsteins’ 35% — 1.9 services instead of 2.4. Health treatment costs came in 23% lower for two-breed crossbreds and 17% lower for three-breed across their lifetimes, with cull value 16% higher.

What does all that do to income over feed cost? Shonka-Martin and colleagues published twice in the Journal of Dairy Science in 2019, and the numbers favour the crossbreds. First-lactation crossbreds ate 2,807 kg (6,189 lbs) of dry matter over 4 to 150 days in milk against the Holsteins’ 2,948 kg (6,499 lbs) — about 5% less — with a further 6.5% reduction later while producing the same fat plus protein. Income over feed cost ran $0.34 to $0.60 higher per cow per day.

Read the inbreeding claim carefully, though. VikingGenetics’ ProCROSS material tells producers there are “no worries about inbreeding,” and for the crossbred animal in front of you that’s essentially right — heterosis does the work. It doesn’t mean the source populations are exempt. “No worries” is marketing shorthand, and the company’s own researchers documented real pressure on those source populations. They documented it in a peer-reviewed journal, which is the opposite of hiding it.

The component question depends entirely on your basis, and this is where a lot of crossbreeding arguments go sideways. On a single 305-day lactation, crossbreds produced less fat plus protein than purebred Holsteins in the Minnesota work — 24 kg (53 lbs) less for Montbéliarde crosses, 29 kg (64 lbs) less for Scandinavian Red. Across a lifetime the picture flips: daily fat plus protein ran 1% higher for two-breed crossbreds and 1% lower for three-breed, because they stuck around long enough to make it up. A rotation is also a multi-year commitment you can’t reverse next season, and crossbred replacements are harder to move if you sell breeding stock.

Then weigh all of it against a replacement market that punishes mistakes. USDA put the U.S. national average for milk cows sold as dairy herd replacements at $3,130 per head in April 2026, up from $2,980 in January and $2,860 a year earlier — 5% on the quarter, 9% on the year. USDA NASS put milk replacement heifer inventory at 3.90 million headon January 1, 2026, down slightly from the prior year. Fixing a genetic misstep costs more than it did two years ago, which argues for care in both directions.

Your Pre-Breeding Action Checklist

  • Pull your inbreeding report in the next 30 days. Run your last three years of sire selections through the Lactanet Inbreeding Calculator or your CDCB-linked mating program and count how many trace to a shared maternal grandsire. A few hours of records work, no new spend, before breeding starts.
  • Run both measures on the same animals, then move your matings to the genomic one. Ablondi’s cows read 7% on pedigree and 16% on genomics, and the depression tracked the bigger figure — 61 kg per point against 44. If you’ve never seen your own two numbers side by side, you don’t know which one your program has been deciding from.
  • If more than two of your last ten sire picks share a maternal grandsire, raise it with your mating consultant. Editorial rule of thumb, not a published standard — no threshold defines “too much overlap” at herd level, so you’re setting a baseline rather than measuring against one.
  • Check whether your TPI cutoff still means what it meant last year. The April 2026 formula change moved bulls up to 125 points without new daughter data. If your program screens on a fixed threshold, some sires dropped off your list on arithmetic alone.
  • Run Pryce’s scaling on your own rolling herd average — 0.4 to 0.6% per point of inbreeding, rather than borrowing an Italian or Dutch figure measured on a different population. At the U.S. rate, a decade costs a 12,000 kg herd 170 to 260 kg per cow per lactation. Multiply your own average by 1.4% to 2.2% to get your version.
  • Ask your supplier for its sourcing split at proof-run specificity, then log the answer or the non-answer. Not proof of anything on its own — but a pattern across several suppliers should shape how much of your lineup comes from one place. Select Sires has published 40% and put an executive on tape saying it; the others haven’t.
  • Before you compare any crossbreeding numbers, establish the basis. Single-lactation and lifetime daily figures point opposite directions on components: 24 to 29 kg less fat plus protein in one lactation, 1% more across a lifetime. Quoting one while meaning the other is the most common way that pitch gets made.
  • Price stillbirth rate as an inbreeding cost, not a calving-management line item. Bjelland’s work puts it in the first column.

Between 2000 and 2009 the industry drove Holstein rate of inbreeding down to 0.37% per generation and pushed Ne up to roughly 135, using optimum contribution selection, expected future inbreeding, and publicly posted relationship values. Those tools worked. Then genomic selection outran them, and they’re still sitting there unused at scale.

The measurement problem is the cheaper of the two fixes. Nobody has to build an origin database or rewrite an index for you to run both numbers on your own females and see which one your matings have been following. Ablondi’s team needed 84,443 markers and 939 herds to prove it at breed scale. You need an afternoon and a report you’re already paying for. The question is whether you’ll like what it says — and whether you’d breed differently if someone finally published where those bulls came from.

Run Your Numbers

Genomic Testing ROI Calculator — This article says pedigree reads low and only genomics shows the real number. The calculator prices what testing is worth on your farm, and flags the families where inbreeding risk needs tighter control. Run the conservative scenario first.

Methodology note: The 7% and 16% figures in this article’s headline are the pedigree and genomic inbreeding averages for Ablondi’s Italian Holstein population and are not presented as a universal read on any individual herd; the size of the divergence depends on pedigree completeness and recent breeding patterns. Nucleus program descriptions and dates are self-reported or from trade media: Select Sires ART (company history page, January 4, 2024 blog post, and a July 2025 Select Sires podcast interview with ART program manager Mark Kerndt, in which the roughly 40% figure refers to bulls sampled each year); PEAK (2012 program start, though official as PEAK in 2018); Semex Progenesis (company pages, founded 2013); ABS Global (company history, first commercial units 2015). USDA-AIPL published the first official U.S. genomic evaluations in January 2009; CDCB assumed responsibility for U.S. genetic evaluations in 2013. Only Select Sires has published a proportional sourcing split. 

Stud-share counts are The Bullvine’s own independent count of the Holstein Association USA Top 200 TPI Genomic Young Bulls list, April 2026 (85% genomic reliability minimum), drawn from the CDCB run published April 7, 2026, with stud assignment by the official NAAB marketing codes table. The April run is cited rather than a later one because Holstein Association USA implemented a TPI formula change in that run, moving production weights from 19% PTA Protein and 19% PTA Fat to 24% PTA Protein and 14% PTA Fat; individual bulls moved up to 125 TPI points with no new daughter data, so April 2026 shares are not comparable to earlier runs and represent the first post-change snapshot. An August 2026 list has since been published and current shares will differ. United Sires, LLC (NAAB 596/796) registered with NAAB on February 8, 2024. Blondin Sires Inc. (NAAB 799, St. Placide, Quebec) is owned by Simon Lalande, Dann Brady and Nicolas Lalande. 

Italian figures: Ablondi, M., A. Summer, G. Stocco, R. Finocchiaro, J.-T. van Kaam, M. Cassandro, C. Dadousis, A. Sabbioni and C. Cipolat-Gotet, Journal of Animal Science 101:skad382 (2023), doi:10.1093/jas/skad382 — 27,735 cows, 939 herds, 84,443 autosomal SNPs, 24 generations of pedigree depth. The 2.35% five-year rate is drawn from Ablondi et al. (2022) and the +415 kg realized progress from ANAFIBJ (2023). Three co-authors are affiliated with ANAFIBJ, which supplied the data and part-funded the work; the authors declare no conflicts of interest. The Parmigiano Reggiano comparison appears in Results and Discussion, using 24-month-aged wheel prices per CLAL (2022), and reflects the difference between the most and least inbred 5% of cows — not a typical per-cow loss. USD and CAD equivalents are editorial conversions at approximately €1 = $1.16 USD and €1 = $1.60 CAD as of September 3, 2026; because the underlying euro figure derives from 2022 cheese prices, the conversion is indicative of present-day value rather than a 2022 dollar equivalent.

Other depression studies use different lactation bases and are not directly interchangeable. Doekes, H.P. et al., Genetics Selection Evolution 51:54 (2019), doi:10.1186/s12711-019-0497-z. Bjelland, D.W., K.A. Weigel, N. Vukasinovic and J.D. Nkrumah, Journal of Dairy Science 96(7):4697–4706 (2013), doi:10.3168/jds.2012-6435 — 205-day, not 305-day. Pryce, J.E., M. Haile-Mariam, M.E. Goddard and B.J. Hayes, Genetics Selection Evolution 46:71 (2014), doi:10.1186/s12711-014-0071-7.

North American population figures: Makanjuola, B.O. et al., Journal of Dairy Science (2020), doi:10.3168/jds.2019-18013 — 676,594 Holstein and 304,676 Jersey pedigree records from the Canadian Dairy Network. Genomic-era Ne is derived from the paper’s reported 1.19–2.06% rate of inbreeding per generation using Ne = 1/(2ΔF), which is why it is reported here as a range rather than a point estimate; an earlier ADSA abstract from the same group reported 30 to 48, differing from the published paper. FAO critical Ne of 50 cited therein (FAO, 2015). Nordic figures: Tenhunen, S., J.R. Thomasen, L.P. Sørensen, P. Berg and M. Kargo, Journal of Dairy Science 107(8):5897–5912 (2024), doi:10.3168/jds.2023-24553; four of five authors are affiliated with VikingGenetics, and the study is open access and peer-reviewed. Methods and periods differ between Makanjuola and Tenhunen, so comparisons are directional.

Generation interval: Guinan, F.L. et al., Journal of Dairy Science 106(2):1110–1129 (2023), doi:10.3168/jds.2022-22205, with corrigendum; figures are for the sire-of-bulls pathway. García-Ruiz et al., PNAS 113(28) (2016), independently found that interval falling from roughly 7 years to under 2.5 between 1975 and 2015. U.S. inbreeding rates: Steyn, Y. et al., Journal of Dairy Science 105(12):9810–9821 (2022), doi:10.3168/jds.2022-22143, citing CDCB (2021).

Crossbreeding: ten-year University of Minnesota study of high-performance Minnesota herds led by L.B. Hansen with A.R. Hazel and B.J. Heins, published 2019. Single-lactation 305-day and lifetime daily component figures use different bases and are reported separately above. Shonka-Martin, B.N., A.R. Hazel, B.J. Heins and L.B. Hansen, Journal of Dairy Science 102:871–882 (2019), doi:10.3168/jds.2018-15318; and Shonka-Martin, B.N., B.J. Heins and L.B. Hansen, Journal of Dairy Science 102:3661–3673 (2019), doi:10.3168/jds.2018-15682. See also Hazel, A.R., B.J. Heins and L.B. Hansen, Journal of Dairy Science (2020). VikingGenetics’ “no worries about inbreeding” statement is a company marketing claim, not independently verified.

Mating programs: Sun, C., S. Westberry, D. Kendall and D. Castellani (STgenetics, Navasota, TX), ADSA Annual Meeting 2019, Abstract #365 — company-authored, not independently validated. Nucleus modelling: Dekkers and Shook, J. Dairy Sci. 73:1920 (1990). Replacement price: USDA, April 2026, U.S. national average with significant regional variation. Heifer inventory: USDA NASS Cattle, released January 30, 2026.

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Nordic Holstein Coancestry Rose Twentyfold. Your Inbreeding Percentage Won’t Show You.

Nordic Holstein female coancestry climbed from 0.02% to 0.39% per year while pedigree inbreeding looked harmless. Coancestry is next generation’s inbreeding — and your current mating report only shows you one of them.

You’re placing a semen order off the August 2026 GTPI list. You take four bulls, spread across the top ten, figuring that spread covers you. Odds are three of them are sons of Ocd Thorson Ripcord-ET — he sired seven of that top ten, and eight of the top ten for Net Merit $, per The Bullvine’s analysis of that run.

Go one generation further back and it gets tighter, not looser. One great-granddam sits behind seven bulls across those two lists.

A proposal circulating in European trade press argues North American studs should voluntarily set aside index thresholds for roughly 5% of young-sire intake, so European-origin sires get a shot they wouldn’t otherwise get. It’s pointing at something real. But it picks the wrong criterion — and the evidence for that runs through a Nordic population where female coancestry climbed nearly twentyfold across the genomic-selection transition, while the pedigree analysis of the same herd found nothing wrong at all.

Holstein Is Climbing at Twice the Rate of Everything Else

Lactanet’s Inbreeding Update – August 2025, authored by Brian Van Doormaal, put Canadian Holstein heifers born in 2024 at 9.99% average pedigree inbreeding. Highest of the four major breeds — ahead of Jersey at 7.56%, Brown Swiss at 7.10% and Ayrshire at 6.89%.

That 9.99% is up from 9.61% for heifers born in 2023. A 0.38-point jump in a single birth year, against a long-run trend of +0.25%. The climb isn’t just continuing. It steepened.

The rate is the number that should bother you. For females born since 2010, Jersey has averaged +0.11% per year, Ayrshire and Brown Swiss +0.12%, and Holstein +0.25%. Better than double.

Worth noting what that rate does and doesn’t tell you. Canadienne actually carries the highest average inbreeding of any Canadian dairy breed at 10.33% — but its rate of increase since 2010 is the lowest of the group at +0.09% per year. High level, slow climb. Holstein is the reverse. Level tells you where a population has been. Rate tells you where it’s going. Hold that distinction, because the whole argument in this piece turns on a version of it.

Four breeds. One country. One evaluation system. The same genomic tools sitting on the shelf for every one of them. Holstein moving twice as fast as the others. Whatever’s driving that, it isn’t a shortage of foreign germplasm — Jersey and Ayrshire breeders in Canada aren’t running set-asides either. (See our companion breakdown: What 9.99% Inbreeding Costs in a Herd Under 50 Cows.)

The Nordic Result Everybody Quotes

At the 2022 World Congress on Genetics Applied to Livestock Production, Tenhunen, Thomasen, Sørensen, Aamand, Berg and Kargo — Aarhus University, VikingGenetics and Nordic Cattle Genetic Evaluations — presented pedigree analysis on 372,955 Nordic Holstein females born January 2016 through November 2018, traced back an average of 12.4 generations.

One term you need before the table. Effective population size — Ne — is the number of breeding animals a population behaves like genetically. Not how many cows exist; how many genuinely independent lines are contributing. A breed with two million cows can run an Ne under 50. The lower it goes, the faster relatedness accumulates. Future Ne is the same idea projected forward off coancestry: where the population is heading if current mating patterns hold. That second number is the one that matters most here.

Their finding travelled fast, and you can see why:

Birth CohortPeriodΔF (per yr)ΔC (per yr)Generation IntervalNeExpected Future Ne
2007–2010Pre-genomics0.00160.00184.86 yr6859
2011–2014Transition0.00170.00144.08 yr7188
2015–2018Genomic era0.00160.00183.17 yr10487

Annual inbreeding rate flat at 0.0016 before and after. Generation interval collapsing from 4.86 years to 3.17. Effective population size apparently rising by half again, and expected future Ne improving from 59 to 87. The authors reported they could not detect any increase in inbreeding or coancestry, contrary to what had been published for Dutch, French and Canadian Holstein.

Two things to hold onto before you bank that. WCGALP proceedings are conference papers, not peer-reviewed journal articles. And the analysis ran on pedigree data.

Then the Same Team Ran It With Genomics

Tenhunen, Thomasen, Sørensen, Berg and Kargo published a follow-up in the Journal of Dairy Science — 2024 Aug;107(8):5897–5912, doi 10.3168/jds.2023-24553 — covering Nordic Jersey and Nordic Holstein, comparing pedigree and genomic measures of inbreeding and coancestry before and after genomic selection, split across females, bulls and approved AI sires. It’s open access, so you can read it yourself.

Nordic Holstein went the other way.

On the genomic measures, effective population size fell from 54.3 before genomic selection to 42.8 after. Future effective population size, the coancestry-derived figure, fell from 198.8 to 42.7 across the same transition.Coancestry rate rose across every Holstein animal group, with the female population surging from 0.02% to 0.39% per year. Yearly ΔF rose for most groups in both breeds.

Two Instruments, Two Verdicts

Measure (source)Pre-GSPost-GSWhat it says
Pedigree Ne — WCGALP 202268104Up 53% — diversity improving
Pedigree expected future Ne — WCGALP 20225987Up 47% — future looks safe
Genomic Ne — JDS 202454.342.8Down 21% — base narrowing
Genomic future Ne — JDS 2024198.842.7Down 79% — the warning
Female coancestry rate, Holstein — JDS 20240.02%/yr0.39%/yr~20x — next generation’s inbreeding
Genomic future Ne, Nordic Jersey — JDS 202440.757.2Up 41% — same program, opposite result

Read that table across the rows, not down the columns. These are not four readings of one quantity. Pedigree Ne and genomic Ne are computed on different scales and cannot be compared to each other by size — 68 is not “bigger than” 54.3 in any meaningful sense. What’s comparable is the arrow. The pedigree instrument said the population was opening up. The genomic instrument, run by the same team on the same population, said it was closing down. Row two and row four are the same conceptual quantity — where the population is headed — measured two ways, pointing opposite directions.

Nordic Jersey, measured the same way over the same transition, went the opposite direction from Holstein: Ne essentially stable, and future Ne improving from 40.7 to 57.2. Same program, same country, same analysis — different outcome by breed.

The authors’ summary is blunt. Genomic methods detect differences between populations and changes in ΔF and ΔC more efficiently than pedigree methods. Genomic selection produced positive coancestry outcomes in Jersey and the opposite in Holstein. Nordic Holstein “faces more pressing concerns,” and the findings “underscore the necessity of genomic control of inbreeding and coancestry with strategic changes to the Nordic breeding schemes.”

So Which Analysis Do You Believe?

The genomic one. That’s not a close call, and it’s the position the research team itself takes.

Be precise about what disagrees here. The two studies don’t offer competing readings of a single number — they used different instruments on the same population and reached opposite conclusions about the direction of travel. Pedigree analysis found diversity holding or improving. Genomic analysis found it deteriorating. One of those instruments missed something.

Pedigree inbreeding measures expected relatedness from recorded ancestry. Genomic measures capture what actually got transmitted — including relatedness that pedigree can’t see because it predates the recorded generations or runs through paths the herdbook doesn’t connect. When the two disagree, the genome is the ground truth and the pedigree is the estimate.

There’s real signal in the 2022 work that survives, and the 2024 paper confirms part of it. Generation interval genuinely fell — across every animal cohort in both breeds, more sharply in males than females. Bulls and AI sires in both breeds showed reduced generational ΔF. Nordic Holstein AI sires were the single group showing a slight decrease in yearly ΔF. The male side of the program did something right.

The Female Side Is Where It Came Apart

Female groups in both breeds showed a negative Ne trend while males were neutral or positive, and Holstein female coancestry climbed nearly twentyfold. A shorter generation interval spun the flywheel faster without widening the base underneath it.

Why the female side specifically? The 2024 paper doesn’t assign a cause, so what follows is inference from how the program is built — but the timing is hard to ignore.

Nordic genomic selection reached full implementation in 2014, the year females were included in the reference populations. Once you can rank a heifer at birth, the elite female pool stops being defined by proven performance across a lactation and starts being defined by a score available on day one. The same top-ranked heifers then get used, repeatedly, as donors. VikingGenetics program material published in 2017 described roughly 10,000 genomic tests on females annually, about 450 heifers contracted for flushing, and some 4,000 embryos produced per year. Its embryo program moves top heifer candidates to a donor station at five to eight months old for flushing or IVF. The stated aim is as many offspring as possible from the top NTM animals.

Note where 2017 sits. It falls inside the 2015–2018 birth cohort the Nordic analysis treats as the genomic era — so those figures describe the program during exactly the window where the coancestry rise shows up.

That is a deliberate and effective design for genetic gain. It is also, by construction, a narrowing of the female base — more calves from fewer dams, selected earlier, on a score that correlates strongly across close relatives. The male side of the program spread its risk across a wider set of sires. The female side concentrated.

One honest caveat in Nordic Holstein’s favour: the authors note ΔF there remains modest compared with what’s been observed in other Holstein populations. Nordic isn’t the cautionary tale. It’s the well-run program whose coancestry problem only became visible when the better instrument came out.

What the Authors Say to Do About It

This is the part that matters for your tank. The 2024 paper’s own conclusion points at mating strategy: analysis of the coancestry data hints at the potential to decrease future inbreeding through informed mating.

In practice that means the coancestry figures identify which pairings will compound relatedness before you make them, rather than showing up as an inbreeding coefficient on a calf that’s already on the ground.

Not passports. Measured relatedness. That’s a selection lever, and it’s the one a herd actually controls — which is a different thing from the supply question the set-aside raises. Hold that distinction; it matters in a minute.

Which is exactly what a Guelph-led team built the tool for.

Somebody Already Built That Tool

Makanjuola, Obari, Condello, Miglior, Maltecca, Cole, Schenkel and Baes — through the Centre for Genomic Improvement of Livestock at Guelph, with Lactanet, NC State, CDCB and Florida — published in the Interbull journal on November 17, 2025.

They pulled Lactanet data: 168,995 genotyped animals, a 616,258-animal pedigree, 8,491 bulls born 2000–2023, and 131,139 cows born 2010–2024. Pedigree completeness above 99%, maximum depth 30 generations. The reference population was active cows and heifers in milk recording with no left-herd date as of the April 2024 test day.

Then they measured how much DNA each bull actually shares with that live Canadian cow population — expected from pedigree (R-value), and realized from genotypes (GR-value).

Across all bulls, R-value ran from 9.3% to 26.5%. GR-value ran from 12.9% to 40.8%. Bulls with United States registration codes came in highest at 20.8% and 30.4%. Bulls registered in the Czech Republic came in lowest, at 17.1% and 24.3%.

Doesn’t the Czech Result Prove the European Case?

Give the set-aside argument its due first, because it raises something a relationship-value screen cannot answer on its own.

Funnel width versus screening precision. You can’t run a GR value on a bull nobody collected. Screening works on the population that already exists — bulls that got sampled, genotyped and entered into an accessible database. The set-aside is about who gets into that population in the first place. Those are two levers on two different parts of the pipeline, and a narrow funnel upstream caps what any downstream screen can possibly find. Widen the funnel and you widen the range of relationship values available to select from. The two are complementary, not competing.

Grant all of that, and the criterion still doesn’t hold.

Read what the Guelph authors actually recommend: select sires with low average relationship values to a defined reference population, as a mating strategy, to reduce or hold inbreeding at acceptable levels while preserving genetic diversity. That’s a relationship-value screen, not a passport screen — and the same logic applies to what you let into the funnel as to what you pick out of it.

The Czech figure isn’t a case for Czech genetics. A Czech-registered bull ranks low here because of how Canadian breeding history ran, not because of where he was born. Run the same math against a different national herd and the order shifts.

And look at the size of the gaps. Between the highest and lowest national groups: about 6 points on the genomic measure. Between individual bulls: nearly 28. The spread inside the bull population is roughly four and a half times the spread between countries. So if you’re going to reserve intake slots — and there’s a real argument you should — reserve them for bulls with low measured relatedness to your reference population, whatever their registration code. A 5% carve-out sorted by passport captures the 6-point axis and leaves the 28-point axis untouched. (Related reading: Genomic Future Inbreeding Shows Relatedness That Pedigree Misses.)

One catch: what the Guelph team published is a method, not a product. Someone still has to run it against your reference population and put the number in front of you. Ask your supplier whether they can — and notice what the answer tells you.

The Number That Isn’t Published Anywhere

The set-aside argument runs on the European-origin bulls that worked. What it doesn’t include is how many got sampled in North America over the same decades and never made a second lineup.

Studs have modelled sampling attrition since at least 1992, when Lohuis published on probability of success and predicted returns for progeny-test sires in the Journal of Dairy Science. Without that denominator broken out by origin, the case rests entirely on the sires that succeeded. Any group of bulls looks strong when only the successes get counted.

Industry Transparency Note: A review of public materials from CDCB, Lactanet, Holstein Association USA and NAAB found no published sire-sampling attrition data broken out by country of origin. That data is held by individual AI companies rather than evaluation bodies, and no association standard or regulatory requirement currently calls for it to be published.

Try to Price the Set-Aside

Meyer et al. (Journal of Dairy Science, 2001) described 1990s U.S. conditions with over 600 new young sires available annually. Five percent of that 1990s figure is about 30 bulls a year across the whole industry — and genomic-era intake is almost certainly lower than 600, which would shrink the number further.

Thirty bulls is small and checkable. It’s also where the math stops. No current published figure exists for genomic-era sampling volume, and no stud has published a per-bull cost breakdown running from acquisition through housing, genomic testing, collection and marketing to a usable proof.

Meanwhile the spread from Net Merit $ #1 to #10 in the August 2026 run is $47 per animal — roughly $4,700 across 100 daughters — per The Bullvine’s analysis of that run. That’s a PTA differential, not money in the tank, but it’s a number the industry can put a decimal on. What it costs to widen the funnel by thirty bulls isn’t published anywhere. (Full run detail: The August 2026 Lists, Bull by Bull.)

The One Historical Case Worth Citing

Carol Prelude Mtoto gets invoked constantly as proof that European testing unlocks genetics North America would have missed. His sire line traces to Ronnybrook Prelude, a Canadian-registered sire confirmed in Holstein Canada’s animal information records. Mtoto’s Italian proving history appears in two independent published accounts rather than a primary herdbook entry, so treat the detail accordingly.

What the case actually shows is a North American pedigree that got its shot in Italy. That’s a story about somebody widening the funnel for an unproven pedigree — which is the set-aside argument’s strongest instinct, and it has nothing to do with where the genetics came from. One case isn’t a pattern either way, and building a procurement policy on it would repeat the same shortcut. (Background: The Full Mtoto Story.)

One Study Nobody Reopened

Powell, Sanders and Norman at USDA’s Animal Improvement Programs Laboratory examined May 2005 Interbull evaluations using Holstein full-brother families — 24,611 bulls for yield traits (Journal of Dairy Science, July 2008, 91(7):2885–92). Bulls from Australia, Germany, Great Britain and Japan showed greater EBV for milk yield than their own full brothers evaluated in the United States, on all countries’ scales. Causes were reported as unknown, and eighteen years on, nobody has replicated it with genomic-era data.

Nilforooshan’s 2022 JDS invited review found that bulls proven in more than one country are “highly selected and a biased representation of the national sire populations.” Sallam et al. (2022) found MACE may not fully account for genotype-by-environment interactions. Neither quantifies direction or magnitude for sires entering North America.

Tie that back to your order. A foreign proof can read differently on a North American scale for reasons that have nothing to do with the bull’s genes — selection bias in which bulls got exported and evaluated abroad, genotype-by-environment effects the conversion may not fully absorb, and in the 2008 case something nobody has identified in eighteen years of trying. None of that makes foreign-tested bulls better or worse than domestic ones. It makes their index numbers harder to read with confidence. Which is one more argument for buying on measured relatedness to your own cow population, where the number means the same thing no matter which country ran the evaluation. An import stamp is not a genetic outcross, and a foreign index is not a like-for-like comparison.

What This Means for Your Operation

What you’re buying onThe numberWhat it actually means
Top 10 GTPI bulls sired by Ocd Thorson Ripcord-ET7 of 10Four picks off the list, three likely half-brothers
Top 10 Net Merit $ bulls sired by Ripcord8 of 10Switching index does not switch families
Bulls tracing to one shared great-granddam7 across both listsOne generation back it tightens, not loosens
Net Merit $ spread, #1 to #10$47/animal (~$4,700 per 100 daughters)A PTA differential, not cash in the tank
Reliability: genomic young sires vs daughter-proven65–82% vs 84–99%On contract heifers, reliability is the decision
Canadian Holstein average inbreeding, 2023 → 2024 heifers9.61% → 9.99% (+0.38 pt in one year)Steeper than its own +0.25%/yr long-run rate

Trust genomic relatedness over pedigree relatedness

Nordic Holstein’s pedigree analysis reported no deterioration. The genomic analysis, from the same team, put future Ne at 42.7 after genomic selection. If your herd manages diversity off pedigree inbreeding alone, you’re reading the estimate instead of the measurement.

Audit your sire-of-sons base

Pull the last twelve months of semen invoices. List sire and maternal grandsire for every straw, then count the distinct sires of sons. Seven of ten top GTPI bulls in the August 2026 run traced to one sire — buy four off that list without checking and the odds say three are half-brothers. Vendor diversity is not family diversity.

Ask for genomic relationship values

Request GR values benchmarked against your national or herd reference base rather than trusting outcross claims on a catalogue page. Individual bulls in the Guelph study spanned 12.9% to 40.8% — roughly four and a half times the spread between the highest and lowest national registration groups. The method exists. Ask who’s willing to run it.

Ask about the funnel too, not just the screen

Screening only works on bulls somebody collected. If your supplier’s young-sire intake is narrow, no amount of GR filtering on the survivors fixes it. Ask how many bulls enter their program each year and how many distinct families those bulls come from — then ask what the relationship-value spread looks like across that intake.

Watch your own donor concentration

The Nordic female result is a warning that applies at herd level. If you’re flushing or aspirating, count how many distinct dams produced your last two calf crops. Genomic pre-selection makes it easy to keep going back to the same three heifers because the score says so. That’s how a female base narrows without anyone deciding to narrow it.

Don’t mistake a shorter generation interval for progress

Nordic Holstein cut generation interval across every cohort and still lost ground on coancestry, because the female side kept drawing on related lines. Faster turnover only expands the effective population if the parents come from genuinely different families.

Watch coancestry, not just inbreeding

Nordic Holstein female coancestry went from 0.02% to 0.39% per year while inbreeding stayed modest. Coancestry is next generation’s inbreeding. Your current inbreeding percentage won’t warn you.

Treat reliability as the decision on contract heifers

Genomic bulls in the August 2026 run carried 65–82% reliability against 84–99% for daughter-proven sires. On heifers you’re committing, that spread is the choice, not the index.

Don’t compare indexes across systems

TPI, LPI, NVI, RZG and gPFT are non-comparable without an explicit MACE-conversion caveat — and treat any foreign-tested proof as carrying an open country-of-testing question, because the 2008 full-brother finding was never revisited.

Key Takeaways

  • If you’re managing diversity off pedigree inbreeding alone, you’re using the instrument that found nothing wrong with Nordic Holstein. In the next 30 days, run the invoice audit above and ask your supplier for genomic relationship values against your national reference base.
  • Funnel width and screening precision are different problems, and you need both. A narrow intake caps what any screen can find; a wide intake sorted by the wrong criterion wastes the width.
  • Canadian Holstein went from 9.61% to 9.99% average inbreeding in one birth year — a steeper jump than its own +0.25% long-run rate, and better than double the pace of Jersey, Ayrshire and Brown Swiss.
  • Same program, same country, same genomic analysis: Nordic Jersey’s future Ne improved from 40.7 to 57.2 across the genomic-selection transition while Holstein’s fell from 198.8 to 42.7. This isn’t an argument about genomics as a technology — it’s about how a breed’s female base gets used.
  • If one bull sits at 30.4% genomic relatedness to your reference population and another at 24.3%, the second one is your outcross — whatever flag is on the catalogue page.
  • If your coancestry rate is climbing while your inbreeding percentage looks stable, that’s the Nordic Holstein pattern, and it resolves into inbreeding one generation later.
  • Ask your stud how many foreign-origin sires they’ve sampled and how many made a second lineup. Whether that number is even available is worth knowing before your next order.

Interbull got built because fragmentation carried a cost the whole industry felt. InterGenomics got built because accuracy had a shared incentive. CDCB publishes its base-change methodology because everybody’s numbers depend on it. Sire attrition data sits at a different level — held by individual companies, with no standard calling for it. Meanwhile the Nordic team went back with a better instrument and published a result that undercut its own earlier finding, and a Guelph-led group has already built the measurement that answers the selection half of the question — and tells you what a wider funnel should be screening for. So the question isn’t whether North America needs European semen. It’s whether anybody’s going to ask their supplier for a relationship value — and whether they’ll get one. What’s yours?

Genetic evaluation figures are from the August 2026 CDCB/Holstein Association USA run. Relationship values are from Makanjuola et al., Interbull journal, November 17, 2025, calculated against the active Canadian cow population as of the April 2024 test day. Inbreeding figures are Canadian, from Lactanet’s Inbreeding Update – August 2025 (Brian Van Doormaal), covering heifers born in 2023 and 2024. Nordic pedigree figures are from Tenhunen et al., WCGALP 2022 conference proceedings; Nordic genomic figures are from Tenhunen, Thomasen, Sørensen, Berg and Kargo, Journal of Dairy Science 2024;107(8):5897–5912, doi 10.3168/jds.2023-24553, open access — all Nordic Ne and future Ne values quoted from that paper are genomic measures compared before and after the genomic-selection transition. Pedigree and genomic Ne are computed on different scales and are not directly comparable by magnitude. Embryo-program figures are from VikingGenetics program material published in 2017, plus its current embryo program description; the 2024 paper does not assign a cause for the female coancestry trend, and the connection drawn here is The Bullvine’s inference. Both Nordic studies cover Denmark, Finland and Sweden. Analysis and conclusions are The Bullvine’s.

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