Breeding selection in modern dairy cattle has evolved from a one-dimensional approach focused solely on maximizing total milk volume into a multidimensional and highly complex biomathematical discipline that integrates genetic potential, metabolic efficiency, environmental sustainability, animal welfare, and bioeconomic profitability. The aggressive single-trait selection pressure applied in the late twentieth century to increase milk yield pushed cows toward their physiological limits and brought with it negative genetic correlations such as lower reproductive performance, greater susceptibility to metabolic disease, and a shorter productive herd life. Today, environmental pressure from climate change, unpredictable global feed costs, and the agricultural sector's commitment to reducing its carbon footprint have made a fundamental redesign of selection-index architecture unavoidable. This comprehensive article examines global changes in phenotypic data standardization, the restructuring of economic indices, feed-efficiency paradigms, biomarkers of heat tolerance, methodologies for methane-emission measurement, and the theoretical framework behind ssGBLUP models.
Global Paradigm Shift
With the widespread adoption of genomic selection in the United States (2009+), the rate of genetic progress increased by 2 to 3 times. As of April 2025, the emphasis on milk fat in the Net Merit (NM$) index increased from 24.7% to 31.8%, while the emphasis on protein declined from 19.6% to 13.0%. The -$57 per point "weight tax" assigned to Body Weight Composite (BWC) reflects the new era of metabolically efficient cattle.
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Calculate Breeding Score1. Introduction: The Evolution of Selection and Today’s Imperatives
Genetic progress in dairy cattle has undergone a dramatic transformation over the last century. In the early 1900s, selection decisions were based largely on phenotypic observation. From the 1930s onward, the launch of systematic milk-recording programs and the adaptation of Jay L. Lush's quantitative-genetic principles to animal breeding placed selection on a scientific foundation and brought Estimated Breeding Value (EBV) calculations into practice. In the 1970s, the BLUP (Best Linear Unbiased Prediction) methodology developed by Charles R. Henderson revolutionized genetic evaluation accuracy by making it possible to separate environmental effects in a systematic way.
However, the intense single-trait selection pressure applied at the end of the twentieth century produced unintended side effects. While milk yield in Holstein populations approximately tripled between 1960 and 2020, fertility indices declined in parallel, the incidence of metabolic disease increased, and average herd life shortened. These negative genetic correlations forced the industry to move toward balanced, multi-trait selection indices.
In 2009, the formal integration of genomic selection into national evaluations in the United States opened a new era in cattle breeding. Thanks to high-density SNP (Single Nucleotide Polymorphism) genotyping panels, the genetic potential of animals could be predicted with high accuracy while they were still heifers. The generation interval in bulls fell from 6-7 years to roughly 2 years, and the annual rate of genetic progress increased by 2 to 3 times.
Today, selection no longer covers only production and health traits; it also includes next-generation phenotypes such as feed efficiency (Feed Saved — FSAV), heat tolerance, methane efficiency (MEF), and sustainability.
2. Phenotypic Data Standardization and the Recalibration of Lactation Curves
The quality of breeding evaluation depends directly on the accuracy of the phenotypic data used and on how well those data can be standardized. To compare animals fairly for genetic potential, sources of environmental variance such as age, parity, calving season, milking frequency, and previous days open must be isolated mathematically.
2.1 The Transition from 305-ME to 305-AA
Since 1935, the dairy industry had relied on 305-day Mature Equivalent (305-ME) standardization. This system was based on estimating the yield a cow would produce at 61-86 months of mature age. However, three decades of intense genomic selection dramatically changed bovine maturation curves. Modern dairy cattle now reach physiological maturity much earlier, and the yield gap between the first and third or fourth lactation has progressively narrowed.
Transition to the 305-AA Standard (August 2024)
USDA AGIL and CDCB re-analyzed 101.5 million milk, 100.5 million fat, and 81.2 million protein lactation records from 1960-2022. As of August 2024, the 305-ME system was abandoned and replaced by the 305-AA (Average Age) system.
Reference Point Shift
Lactation records are now standardized to an average age of 36 months (the start of second lactation) rather than to a theoretical mature age. The former ME system overstated records by roughly 10% compared with actual production.
Expanded Climate Regions
Instead of the former three geographic regions, the new model defines five specific climate regions based on average climate scores. The share of seasonal variance within total variance has narrowed with the spread of closed-housing technologies.
Breed-Specific Coefficients
Instead of Holstein-dominated general coefficients, specific coefficients are now calculated for each breed's developmental biology, including Jersey, Brown Swiss, Ayrshire, and others.
2.2 Effects of the 305-AA Transition on PTA and Economic Indices
| Breed | PTA Milk Change | PTA Fat/Protein | Impact on NM$ |
|---|---|---|---|
| Holstein | Upward recalibration | Upward | +$10 to +$15 |
| Jersey | Decrease of about 100 lb | Decrease of about 6 lb (fat + protein) | -$50 to -$70 |
| Brown Swiss | Minimal change | Near zero | Stable |
| Ayrshire / Guernsey | Minimal change | Minor fluctuation | Stable |
Important Warning
These results prove that updating phenotypic standardization models can have a seismic impact on genetic rankings. It is therefore critical for breeders to know which standardization system was used when interpreting bull-catalog values.
3. Genetic-Base Updates and the Integration of New Phenotypic Traits
The genetic base, which serves as the reference point for genetic evaluations, is updated periodically. The genetic base is the baseline at which the average PTA values of cows born in a given year are set to zero. In most populations, this base is shifted forward every five years.
3.1 April 2025 Genetic-Base Update
Why Is the Base Updated?
- Genetic progress is continuous, so the gap between the old base and new animals keeps widening
- PTA values become inflated over time, and a new base corrects this inflation
- It restores meaningful comparison with the current population
- Standard deviations for milk, fat, protein, and SCS are recalculated by breed
Points Requiring Attention
- The PTA of the same bull declines mathematically after a base change; this is not a genetic loss
- Selection thresholds derived from older references become incompatible
- Different countries use different base years, so international comparisons require caution
- Holstein PTA Milk: about -750 lb; PTA Fat: about -45 lb (mathematical shift)
3.2 New Phenotypic Trait: Milking Speed (MSPD)
Advances in data-collection technologies are making it possible to bring new phenotypes into breeding decisions. As of August 2025, the Milking Speed (MSPD) trait for Holsteins has been completely redesigned. The new MSPD uses objective data from in-line sensor technologies in milking systems instead of classifier-based subjective scoring. Based on an average flow rate of 7 lb (3.18 kg) per minute, this phenotype has become a critical criterion for modern milking-parlor efficiency and for the selection of cows suited to automatic milking systems (AMS).
4. Effects of Local Environmental Factors and Morphological Traits on Milk Yield
In addition to global standardization models, understanding the local dynamics of genotype × environment interactions (GxE) is essential for regional adaptation and productivity. Regional studies examining how environmental variance operates at the local level and how specific physical conformation traits relate to production provide important feedback when applying global indices to local populations.
4.1 Studies at the Bahri Dagdas International Agricultural Research Institute
In Turkey, long-term studies conducted on Brown Swiss cattle by Mehmet Colak and his research team at the Bahri Dagdas International Agricultural Research Institute provide noteworthy findings in this context. In analyses of lactation records from Brown Swiss cattle raised under institute conditions between 1987 and 1999, the heritability of 305-day milk yield was estimated at 0.23 for first lactation and 0.19 across all lactations (Tilki, Colak & Sari, 2009).
Bahri Dagdas Research Findings
The finding that 305-day milk yield heritability in Brown Swiss cattle was h² = 0.23 in first lactation and h² = 0.19 across all lactations confirms the dominant influence of environmental factors on milk production. These moderate heritability estimates provide a scientific explanation for why phenotypic correction factors are critical in breeding evaluation.
4.2 The Relationship Between Udder Morphology and Milk Production
The same research group also examined in detail the relationships between udder morphology and phenotypic milk production. In studies investigating the effect of teat shape on milk yield and milking characteristics, teat shape was classified as cylindrical, funnel-shaped, and bottle-shaped (Tilki, Inal, Colak & Garip, 2005).
| Teat Shape | Average 305-Day Milk Yield (kg) | Statistical Interpretation |
|---|---|---|
| Cylindrical | 3,156 | Statistically higher with a significant difference (P < 0.05) |
| Funnel-shaped | 3,169 | |
| Bottle-shaped | 2,377 | Significantly lower yield |
In another study, 400 teat canals were examined by ultrasonography, and 20% were found to have a crooked course while 80% followed a linear course. This anatomical variation did not produce a statistically significant difference in final milk yield. Measurements such as front and rear teat length, teat diameter, and post-milking udder height showed statistically significant changes across lactation order (P < 0.001).
How Morphological Studies Support Selection Indices
Such local morphological studies provide the biological rationale for integrating functional type traits such as Udder Composite (UDC) into modern selection indices with substantial economic weights. Udder structure influences not only milking speed, but also mastitis resistance and productive life in the herd.
4.3 Environmental Correction Factors: Comparing Turkish and Global Literature
Environmental correction coefficients derived from Bahri Dagdas studies show high agreement (90%+) with current global literature, including CDCB 305-AA, ICAR 2024, and JDS 2025:
Parity Coefficient (Base = 3rd-4th lactation = 1.00)
| Lactation | Coefficient | Turkey (DSYMB) | Global (CDCB/ICAR) |
|---|---|---|---|
| 1st lactation | ×1.13 (1.12-1.15) | Average ×1.13 | ×1.12-1.15 ✓ |
| 2nd lactation | ×1.06 (1.05-1.08) | Consistent | ×1.05-1.08 ✓ |
| 3rd-4th lactation | ×1.00 | Base | Base ✓ |
| 5th+ lactation | ×0.94 (0.92-0.96) | ×0.93-0.96 | ×0.92-0.95 ✓ |
Age-at-Calving Coefficient (Base = 37-48 months = 1.00)
| Age Group | Coefficient | Interpretation | Global Alignment |
|---|---|---|---|
| <30 months | ×1.18 (1.15-1.20) | Largest correction | USDA: ×1.17-1.20 ✓ |
| 31-36 months | ×1.10 (1.08-1.12) | Moderate correction | ✓ |
| 37-48 months | ×1.00 | Reference | CDCB 36-month base ✓ |
| 49-60 months | ×0.97 | Slight decline | ✓ |
| >60 months | ×0.93 (0.92-0.95) | Longevity effect | ×0.92-0.95 ✓ |
Season Coefficient (Base = Spring = 1.00)
| Season | Coefficient | Interpretation | Global Alignment |
|---|---|---|---|
| Winter (Dec-Feb) | ×1.08 (1.05-1.10) | Highest yield | ×1.07-1.10 ✓ |
| Spring (Mar-May) | ×1.00 | Reference | Base ✓ |
| Summer (Jun-Aug) | ×0.93 (0.90-0.95) | Heat stress | 7-10% decline ✓ |
| Autumn (Sep-Nov) | ×1.01 (0.98-1.02) | Close to spring | ×1.00-1.02 ✓ |
Adjusted 305-Day Milk Formula
Multiplicative correction formula used to standardize the animal's raw yield:
Adjusted 305-AA Milk (kg) = Raw Milk × Parity × Age at Calving × Season × THI Correction
The VetKriter Breeding Potential Score calculator applies this adjustment automatically.
5. Restructuring Economic Selection Indices: The NM$ 2025 Revision
The main mechanism through which genotypic data become actionable in the field is the economic selection index. In the United States, the principal national index is the Lifetime Net Merit Index (NM$), which estimates the net profit, in U.S. dollars, generated over the lifetime of the average daughter of a dairy animal. The NM$ formula includes approximately 40 economically relevant traits:
Core NM$ Formula
NM$ = a' × u
a = the economic-value vector for each trait (marginal contribution to profit); u = the animal's PTA vector. Some traits, such as SCS, are included after normalization.
5.1 The Rise of Butterfat Value and the Relative Decline of Protein
In the 2025 economic model, the fat price per pound for Class III milk was increased from $2.10 to $2.80, while the protein price was kept fixed at $2.60:
Butterfat
24.7% → 31.8%
The most heavily weighted trait
Protein
19.6% → 13.0%
Substantial decline
5.2 Livability Versus Productive Life
| Economic Parameter | Previous Value | 2025 Update | Change |
|---|---|---|---|
| Cull-cow price (lb) | $0.60 | $0.90 | +50% |
| Heifer calf value | $200 | $400 | +100% |
| Cow death-loss cost | $1,800 | $2,038 | +13% |
Because the financial gap between a cow going to slaughter (economic recovery) and a cow dying on the farm (total loss) has widened, Cow Livability (LIV) and Heifer Livability (HLIV) now carry much greater weight. The overall emphasis on Productive Life (PL) has been reduced from 11.0% to 8.0%.
5.3 The Weight Tax
The old assumption that "a bigger cow produces more milk" has given way, in the genomic era, to the concept of metabolic efficiency. Increasing feed cost from $0.11 to $0.12 per pound, maintenance dry-matter requirement per lactation from 4.50 to 5.50 lb, and heifer-rearing cost from $0.75 to $0.85/lb has made large cows more expensive to maintain. A +1.0-point increase in BWC corresponds to -$57 in lifetime NM$, expressed economically as a "weight tax."
5.4 Innovations in Fertility Algorithms
The spread of sexed semen and beef-on-dairy crossbreeding has fundamentally changed reproductive economics. The 2025 model abandons the historical assumption of 100% conventional dairy semen and instead incorporates the use of 60% sexed dairy semen (at $25 per dose) and 40% conventional beef semen (at $10 per dose).
Profitability by Lactation
1st lactation
-$120
Loss due to high rearing cost
2nd lactation
+$151
Beginning of profitability
3rd lactation
+$209
Peak profitability
These figures explain mathematically why fertility and health are indispensable for profitability.
5.5 Relative NM$ Economic Weights by Breed (April 2025)
| Trait / Category | Holstein (%) | Jersey (%) | Brown Swiss (%) | Ayrshire (%) |
|---|---|---|---|---|
| Fat | 24.7 | 29.9 | 26.9 | 30.2 |
| Protein | 11.4 | 15.2 | 14.3 | 15.2 |
| Milk Volume | 2.9 | 3.4 | 3.3 | 3.7 |
| Productive Life (PL) | 12.3 | 15.8 | 18.0 | 16.9 |
| Residual Feed Intake (RFI) | -14.2 | 0 | 0 | 0 |
| Body Weight Composite (BWC) | -11.2 | -13.0 | -13.0 | -14.4 |
| Cow Livability (LIV) | 5.9 | 6.0 | 6.6 | 5.1 |
| Udder Composite (UDC) | 1.2 | 1.0 | 1.3 | 2.0 |
| Somatic Cell Score (SCS) | -2.6 | -2.6 | -3.3 | -3.6 |
| Daughter Pregnancy Rate (DPR) | 2.6 | 3.6 | 3.4 | 3.5 |
Table data: USDA ARR NM$9 2025 revision. RFI data are currently available only for Holsteins; because they are not measured in other breeds, the weights of other traits increase proportionally.
NM$ Variants
Cheese Merit (CM$): Negative weight on milk volume, high value on protein. Fluid Merit (FM$): Protein value set to zero, with emphasis only on volume and fat. Grazing Merit (GM$): Assigns roughly 2.5 times the weight of standard indices to fertility traits such as DPR and CCR.
6. Feed-Efficiency Paradigms: Feed Saved (FSAV) and RFI
In dairy farming, more than half of total operating costs come directly from feed expenses. The Feed Saved (FSAV) trait, incorporated into genetic evaluations by CDCB in December 2020, is one of the central components of the "Income Over Feed Cost (IOFC)" optimization paradigm.
6.1 Definitions of FSAV and RFI
FSAV is a composite index synthesized from two components: Body Weight Composite (BWC) and Residual Feed Intake (RFI).
What Is RFI?
RFI is the difference between a cow's actual feed intake and the expected feed intake calculated from body size, growth, milk production, and body condition. Negative RFI indicates a metabolically superior animal that consumes less feed than expected. Its heritability is h² ≈ 0.19, higher than that of somatic cell score or pregnancy rate.
Example of an FSAV PTA
Daughters of a Holstein bull with an FSAV PTA of +200 consume, on average, 200 lb (~90 kg) less dry matter per lactation than their peers. By definition, the genetic correlation of RFI with production traits is near zero, which means metabolically efficient cows can be selected without sacrificing milk yield.
Link to Environmental Sustainability
It has been scientifically demonstrated that cows consuming less feed also produce less manure and, consequently, lower methane emissions. Feed efficiency has therefore become a key element of environmental sustainability. To avoid double counting in the NM$ formula, the economic weights of BWC and RFI are integrated in a balancing way (in 2025, BWC -11.0%, RFI -6.8%).
7. Genomic Adaptation to the Climate Crisis: Heat Tolerance
Rising environmental temperatures driven by global climate change are among the greatest biological risks threatening dairy production. Modeling studies project that uncontrolled heat stress could impose an annual economic burden of US$30 billion on the global dairy industry by 2050.
7.1 The Physiological Cascade of Heat Stress
THI (Temperature-Humidity Index) Formula
THI = (1.8 × Tdb + 32) − [(0.55 − 0.0055 × RH%) × (1.8 × Tdb − 26)]
Tdb = dry-bulb temperature (°C); RH% = relative humidity. Heat stress begins at a THI threshold of approximately 68-72.
Once THI exceeds the threshold, cows redirect blood flow from internal organs toward peripheral tissues in order to dissipate body heat. This results in the following pattern:
| THI Range | Stress Level | DMI Loss | Effect on Milk Yield | Correction Factor |
|---|---|---|---|---|
| ≤67 | No stress | — | Normal | ×1.00 |
| 68-72 | Mild | 0.5-1.0 kg/day | 3-5% decline | ×0.97 |
| 73-77 | Moderate | 1.5-2.5 kg/day | 5-10% decline; SCC ↑ | ×0.93 |
| 78+ | Severe | 2.5-3.75 kg/day | 10-25% decline; embryonic loss ↑ | ×0.88 |
7.2 Reaction-Norm Models and Heat-Tolerance GEBVs
Zoetis and other genetic-research consortia have developed specific heat-tolerance genomic breeding values (GEBVs) by combining millions of test-day records with weather data. These evaluations focus on two traits: resistance to pregnancy-rate decline at increasing THI (CFS_THI) and resistance to milk-yield decline (MILK_THI). Validation studies have clinically confirmed that cows with high CFS_THI and MILK_THI transmitting abilities can maintain rectal temperatures below 39°C even during severe heat waves (≥25°C).
7.3 Molecular Biomarkers and the SLICK Variant
Genome-wide association studies (GWAS) have demonstrated that QTL on chromosomes BTA 3, 6, 14, and 17 are directly associated with heat tolerance. Transcriptomic analyses have shown that, under thermal stress, the expression of lactation-related genes in mammary tissue (α-casein, β-lactoglobulin) declines by 30-40%, while mitochondrial-fusion genes such as Mfn2 are downregulated.
The SLICK Gene: The Most Radical Genomic Discovery
The SLICK haplotype, originally identified in Caribbean Senepol cattle, results from a dominant mutation in the Prolactin Receptor (PRLR) gene. It gives animals a characteristically short, sparse hair coat, maximizing sweat-gland activity and heat dissipation through the skin. Through introgression programs, it has been successfully transferred into high-producing Holstein and Jersey populations. Cows carrying SLICK have been shown to substantially reduce milk-yield collapse during heat waves and to expand their thermoneutral zone.
8. Environmental Sustainability: Genomic Evaluation of Methane Emissions
Ruminant livestock production is responsible for approximately 14.5% of global greenhouse-gas emissions attributable to human activity. A major share of this comes from methane (CH₄), whose global warming potential is 28 times higher than that of CO₂. Methanogenic Archaea in the rumen combine metabolic hydrogen with CO₂ generated during fermentation to synthesize methane. This enteric methane represents an irreversible loss of 4-7% of the gross energy consumed by the animal.
8.1 ICAR Section 20: Framework for Genetic Evaluation of Methane
Through its Section 20 guidelines, the International Committee for Animal Recording (ICAR) has established the global technical framework for standard genetic evaluation of methane emissions in dairy cattle. Scientific studies have shown that methane production (MeP — g/day), methane intensity (MeI — g CH₄/kg milk), and methane yield (MeY — g CH₄/kg DMI) are measurable traits with heritability estimates ranging from h² = 0.12 to 0.35.
8.2 Methodologies for Measuring Methane
| Method | Accuracy | Cost | Data Volume | Use Case |
|---|---|---|---|---|
| Respiration chambers (RC) | Gold standard | Very high | Very low | Research |
| SF₆ tracer technique | High | Moderate | Low | Research / field |
| GreenFeed systems | High | Moderate | Moderate | Field research |
| Sniffer devices (AMS) | Moderate | Low | Very high | Robotic milking systems |
| Laser methane detectors | Variable | Low | High | Field screening |
8.3 Milk MIR Spectroscopy: A Transformative Phenotypic Proxy
Because it is impossible to test millions of cows directly with measurement devices on a national scale, milk mid-infrared (MIR) spectroscopy has become the most important phenotypic proxy. During routine milk analysis, MIR devices read the absorption patterns of infrared light passing through milk samples and generate detailed spectral data. Partial least squares (PLS) regression models and artificial-intelligence algorithms combine these MIR spectra with milk yield, DIM, and parity information to estimate a cow's daily enteric methane production with an accuracy of R² = 0.60-0.70.
8.4 Canada’s Leadership: The Methane Efficiency (MEF) GEBV Model
The first country to integrate this technology into a national breeding program was Canada (the Lactanet consortium). After processing more than 13 million milk MIR records, Lactanet officially released a Methane Efficiency (MEF) GEBV for Holsteins in April 2023.
Technical Details of the MEF Model
- Software: MiX99 — a single-step four-trait animal model (Milk, Fat, Protein, CH₄MIR)
- Critical innovation: Recursive genetic linear-regression coefficients make methane production genetically independent from milk and component yields (Residual Methane)
- Presentation: Population mean = 100, standard deviation = 5. Values above 100 indicate a genetic background associated with lower methane emissions
- Target: A 20-30% genetic reduction in methane emissions per cow by 2050 without sacrificing milk yield
- Other countries: Australia, Spain, Denmark, and the Netherlands are moving similar indices toward commercial use
9. Herd-Level Sustainability Recording Traits (ICAR Section 22)
As of July 2023, ICAR has published Section 22, the guideline for "Sustainability Recording Traits." This framework defines 43 standardized traits that organizations can use to build their own sustainability indices. To eliminate climatic and seasonal bias statistically, nearly all of these traits are calculated over a 365-day rolling window.
| Category | Main Traits |
|---|---|
| Feeding and Production | Energy-corrected milk (ECM), functional BCS ratio, methane emissions, MUN ratios, dry-matter intake |
| Fertility | Calving interval, 56-day non-return rate (NR56), first-service conception rate, days open, visible pregnancy loss |
| Health | Mean SCS, chronic infection rate, dry-period treatment success, fresh-cow infection, proportion of first-test FPR < 1 or > 1.3 |
| Longevity | Average number of lactations, age at culling, lifetime average production, cow mortality within the first 60 days |
| Young Stock | Age at first calving, stillbirth rate, involuntary culling in young stock, respiratory/diarrheal losses |
9.1 Dry-Matter Intake and Methane Prediction Equations
For situations in which collecting actual feed-intake or methane-emission data at the farm level is impossible, ICAR provides comprehensive biomathematical prediction equations. The dry-matter intake estimate based on NASEM (formerly NRC) modeling is:
NASEM DMI Prediction Formula
DMI (kg/day) = [(0.372 × ECM + 0.0968 × CA0.75) × (1 − e−0.192 × (DIM/7 + 3.67))] × Parity Correction
CA = body weight (kg); ECM = energy-corrected milk; DIM = days in milk; parity = 0 (primiparous) or 1 (multiparous).
In high-forage scenarios, more complex regression equations based on dietary fiber and fat proportions are used for enteric methane (eCH₄), such as the Escobar-Bahamondes model. These formula sets allow genetic selection to be integrated with farm-management software and make sustainability targets statistically trackable.
10. Advanced Genomic Modeling: The Architecture of Single-Step Genomic BLUP (ssGBLUP)
In modern dairy cattle, single-step genomic BLUP (ssGBLUP) models have become the industry standard for converting phenotypic data into genetic merit (GEBV). The revolutionary feature of this methodology is its ability to combine pedigree records, phenotypic data, and SNP profiles from both genotyped and nongenotyped animals within one massive simultaneous system of equations.
10.1 The H Matrix: Combining Pedigree and Genomic Information
In classical BLUP, the genetic variance-covariance structure is modeled solely by the pedigree-based numerator relationship matrix (A). However, this approach cannot capture the fact that full siblings receive different gene combinations through Mendelian sampling. ssGBLUP merges the pedigree matrix (A) and genomic matrix (G) to create a composite combined relationship matrix (H):
The H⁻¹ Matrix (Henderson Formulation)
H⁻¹ = A⁻¹ + [0, 0; 0, Gw⁻¹ − A22⁻¹]
A⁻¹ = inverse pedigree matrix for the full population; A22⁻¹ = inverse pedigree matrix for the genotyped subset; Gw = blended genomic matrix. Through the pedigree network, the model propagates genomic information both backward and forward to related animals that have not been genotyped.
10.2 Blending and Scaling
The incompatibility between A and G arises because pedigrees typically cover only a few generations, whereas SNP markers reflect mutation history over thousands of years. This mismatch in base population can lead to overprediction in the genomic values of young bulls. The blending operation is:
Blending Formula
Gw = (1 − β) × G + β × A22
β, the blending parameter, is generally chosen in the range of 0.30-0.40. This procedure both guarantees positive definiteness of the matrix and incorporates residual polygenic variation not captured by the SNP panel. Optimization studies show that β = 0.30-0.40 and ω = 0.60 provide a 6-7% net increase in GEBV accuracy.
10.3 Unknown Parent Groups (UPG) and Metafounders
To address problems caused by incomplete pedigree data, Unknown Parent Groups (UPG) and the more advanced concept of Metafounders (MF) are integrated into the H⁻¹ matrix. These concepts account for the genetic level of the generation to which missing ancestors belong, preventing animals with unknown parents from collapsing to zero in the model and distorting the genetic trend. Modern computational software solves multiple traits and data from tens of millions of animals through these matrix structures, forming the statistical foundation of the genomic era.
11. Practical Breeding-Selection Criteria and Next-Generation Traits
After reviewing the theoretical background and global indices, it is necessary to present, in hierarchical order, the concrete criteria that determine a cow's breeding potential at the field level. The table below summarizes the main criteria that should be considered in modern dairy-cattle breeding decisions, in order of priority:
| # | Criterion | Subcomponents | Weight Range | Explanation |
|---|---|---|---|---|
| 1 | Health and Genetic Foundation | SCS, mastitis resistance, EBV for metabolic disease, genomic test result (GTPI/gRZG/NM$) | 25-30% | An unhealthy cow means unsustainable production. If genomic testing is available, reliability increases dramatically. |
| 2 | Adjusted Milk Yield + Composition | 305-AA milk (kg), fat %, protein %, fat + protein total (kg) | 30-35% | Standardized performance after correction factors are applied. Total fat + protein yield is more valuable than raw milk volume. |
| 3 | Reproductive Performance | Calving interval, DPR/CCR, first-service conception, days open | 10-15% | Regular calving means regular revenue. Fertility problems directly affect herd economics. |
| 4 | Conformation and Longevity | UDC, FLC, PL/LIV, BWC | 10-15% | Udder structure, foot and leg soundness, and stayability in the herd. BWC is now selected in a negative direction. |
| 5 | Sustainability and Emerging Traits | FSAV/RFI, methane EBV, heat tolerance, A2A2, κ-casein BB | 5-10% | Forward-looking value. Current data are still limited, but economic importance is increasing. |
11.1 Milk-Protein Genetic Variants: A2A2 and κ-Casein BB
The beta-casein A2A2 genotype means that the β-casein protein in milk contains only the A2 variant. Epidemiological studies have suggested that the A1 variant releases the bioactive peptide beta-casomorphin-7 (BCM-7) during digestion and may be associated with gastrointestinal discomfort in some individuals. The global A2 milk market is expanding rapidly, and cows with the A2A2 genotype can provide a premium-pricing advantage for producers.
The kappa-casein (κ-CN) BB genotype is critically important for cheese manufacture. Compared with milk from AA cows, milk from BB cows has better coagulation properties and increases cheese yield by 5-10%. Given the share of cheese production in Turkey's total milk utilization, selection for κ-CN BB carries clear strategic economic value.
11.2 Integration of Genomic Testing
GTPI (USA)
Total Performance Index. The most widely used aggregate index for Holsteins. It combines milk production, health, fertility, and type traits. GTPI > 3000 indicates top-tier genetic potential.
gRZG (Germany)
Genomischer Relativzuchtwert Gesamt. Mean = 100, SD = 12. gRZG > 130 indicates elite genetics. It places greater weight on health and functional traits.
NM$ (USA)
Net Merit Dollar. Lifetime net economic value. NM$ > 1000 indicates high profitability potential. As of April 2025, the weight assigned to fat increased to 31.8%.
12. Breeding Selection Practice and Institutional Infrastructure in Turkey
In Turkey, dairy-cattle breeding is coordinated by the Central Association of Cattle Breeders of Turkey (DSYMB) and its provincial associations. The milk-recording system captures milk yield, fat, and protein percentages using A4 (monthly recording) and AT (alternate testing) methods. Herdbook records are maintained in accordance with ICAR standards.
12.1 Current Situation and Challenges in Turkey
Strengths
- The DSYMB milk-recording infrastructure is active and expanding
- Strong research institutes such as Bahri Dagdas and Lalahan
- The size of the Holstein population is sufficient for genomic evaluation
- The younger generation of breeders is open to technology
- The number of farms using genomic testing is increasing rapidly
Areas Requiring Improvement
- The national genomic evaluation system is not yet fully operational
- Reference-population development is still in progress
- Infrastructure for collecting feed-efficiency (RFI) and methane data is inadequate
- Record-keeping rates remain low in small and medium-sized farms
- Heat-stress management is critical in the Southeast, Mediterranean, and Aegean regions but is still insufficiently monitored
12.2 Practical Recommendations
A Breeding-Selection Strategy for Turkey
- Keep records: Participate in a milk-recording program without fail. The genetic value of an unrecorded animal cannot be estimated.
- Use correction factors: Do not compare raw milk yield directly; apply adjustments for age, parity, and season.
- Use genomic testing: Especially in elite cows and heifers considered as bull-dam candidates. The cost-benefit ratio is highly favorable.
- Use aggregate indices: Do not select bulls on milk yield alone. Prefer balanced indices such as NM$, gRZG, or TPI.
- Emphasize health and fertility: Prefer bulls with low SCS, high DPR, and long PL.
- Consider heat tolerance: In hot regions with many days above THI > 72, consider bulls with high heat-tolerance transmitting abilities or crossbreeding strategies (for example Jersey or Montbeliarde).
- Give importance to fat and protein: Total fat + protein yield (kg) is more valuable than pure milk volume. If your enterprise is cheese-oriented, prioritize the κ-CN BB genotype.
- Select for A2A2 when relevant: If the A2 milk market is expanding, increase the A2A2 frequency in your herd to capture long-term premium potential.
13. Conclusion and Future Perspective
Breeding selection in dairy cattle is no longer simply a matter of choosing "the bull that produces the most milk." It is now a multidimensional decision process that integrates biological efficiency, environmental sustainability, animal welfare, and economic optimization. The spread of genomic technologies, the update of phenotypic standardization through 305-AA, the restructuring of economic indices such as NM$ 2025, and the rise of advanced statistical models such as ssGBLUP as industry standards are the core pillars of this transformation.
The dairy cow of the future will be an "eco-sustainable" animal that produces milk rich in fat and protein, consumes less feed (negative RFI), emits less methane, resists heat stress, remains healthy, and lives longer. Achieving this goal will only be possible through the integrated application of accurate phenotypic data collection, reliable genomic evaluation, and science-based selection decisions.
Vision for 2030-2040
- Methane emissions: Targeting a 20-30% reduction per cow through genetic selection
- Feed efficiency: RFI data will become part of routine evaluation in all major breeds
- Heat tolerance: SLICK and other thermoregulation genes will be introgressed into dairy populations
- Precision livestock farming: Sensor data on activity, rumination, and body temperature will be integrated into genomic models in real time
- Artificial intelligence: Machine-learning algorithms will improve the accuracy of multi-trait GEBV prediction
- Turkey: Completion of a national reference population and integration of a domestic genomic evaluation system with the global framework
14. References
- Tilki, M., Çolak, M., & Sari, M. (2009). Genetic parameters of 305-day milk yield for Brown Swiss reared in the Bahri Dagdas International Agricultural Research Institute in Turkey. Kafkas Üniversitesi Veteriner Fakültesi Dergisi, 15(5), 801-804.
- Tilki, M., İnal, Ş., Çolak, M., & Garip, M. (2005). Relationships between milk yield and udder measurements in Brown Swiss cows. Turkish Journal of Veterinary & Animal Sciences, 29(1), 75-81.
- Miles, A. M., VanRaden, P. M., & Norman, H. D. (2025). Updated 305-day lactation yields for age, season, milking frequency, and pregnancy by climate region and breed. Journal of Dairy Science, 108(1), 775-789.
- VanRaden, P. M., Cole, J. B., & Parker Gaddis, K. L. (2025). Net Merit as a Measure of Lifetime Profit: 2025 Revision (NM$9). USDA AIP Research Report.
- Rojas de Oliveira, H., Brito, L. F., Lourenco, D. A. L., & Miglior, F. (2024). Genetic evaluation for methane efficiency in dairy cattle using milk mid-infrared spectroscopy. JDS Communications, 5(2), 178-183.
- Dikmen, S., Khan, M. J., Huson, H. J., & Hansen, P. J. (2025). Heat stress abatement in cattle through genetic improvement of heat tolerance. Journal of Animal Science, 103(2), skae378.
- Aguilar, I., Misztal, I., Johnson, D. L., Legarra, A., Tsuruta, S., & Lawlor, T. J. (2010). Hot topic: A unified approach to utilize phenotypic, full pedigree, and genomic information for genetic evaluation of Holstein final score. Journal of Dairy Science, 93(2), 743-752.
- Bermann, M., Lourenco, D., & Misztal, I. (2022). The distribution of inbreeding depression and recovery by genomic regions in single-step genomic BLUP. Italian Journal of Animal Science, 21(1), 611-622.
- Misztal, I., Legarra, A., & Aguilar, I. (2014). Using recursion to compute the inverse of the genomic relationship matrix. Journal of Dairy Science, 97(6), 3943-3952.
- ICAR (2020). Section 20: Enteric Methane Emissions Recording Guidelines. ICAR Guidelines.
- ICAR (2023). Section 22: Sustainability Recording Traits. ICAR Guidelines.
- Wiggans, G. R., Cole, J. B., Hubbard, S. M., & Sonstegard, T. S. (2017). Genomic selection in dairy cattle: The USDA experience. Annual Review of Animal Biosciences, 5, 309-327.
- Pryce, J. E., & Haile-Mariam, M. (2020). Symposium review: Genomic selection for reducing environmental impact and adapting to climate change. Journal of Dairy Science, 103(6), 5366-5375.
- De Haas, Y., Pszczola, M., Soyeurt, H., Wall, E., & Lassen, J. (2017). Invited review: Phenotypes to genetically reduce greenhouse gas emissions in dairying. Journal of Dairy Science, 100(2), 855-870.
- Tempelman, R. J., Spurlock, D. M., Coffey, M., Veerkamp, R. F., Armentano, L. E., ... & VandeHaar, M. J. (2015). Heterogeneity in genetic and nongenetic variation and energy sink relationships for residual feed intake across research stations and countries. Journal of Dairy Science, 98(3), 2013-2026.
- Legarra, A., Aguilar, I., & Misztal, I. (2009). A relationship matrix including full pedigree and genomic information. Journal of Dairy Science, 92(9), 4656-4663.
- DSYMB (2024). Turkey Cattle Breeders Central Association Milk Recording Reports. Ankara.