Understanding Combining Ability in Maize: GCA, SCA & Heterosis

Combining Ability in Maize

Last Updated: November 4, 2025

Estimated reading time: ~6-7 minutes

Word count: 1391

The development of high-yield maize (Zea mays L.) hybrids is a cornerstone of global food security. But how do breeders know which two parent plants will create a superstar offspring? The answer lies in the complex genetic concepts of combining ability, gene action, and heterosis. A 2024 doctoral thesis by Indrajeet Kumar provides a comprehensive framework for understanding these principles, which are essential for both plant breeding and applied zoology, especially when studying plant-pest interactions and crop improvement.

  • Learn the difference between General Combining Ability (GCA) and Specific Combining Ability (SCA).
  • Understand how GCA is linked to additive gene action, while SCA is linked to non-additive (dominance) gene action.
  • Define heterosis (hybrid vigor) and the genetic hypotheses that explain it.
  • Explore why Genotype x Environment (GxE) interaction is a major challenge for breeders.

The Core Concept: Combining Ability (GCA vs. SCA)

In maize breeding, “combining ability” refers to a parent line’s capacity to produce superior (or inferior) offspring when crossed with other lines. Breeders categorize this into two distinct types: General Combining Ability (GCA) and Specific Combining Ability (SCA).

Professor’s Insight: GCA and SCA are the most important predictive tools a breeder has. GCA tells you if a parent is “good” on average, while SCA tells you if a specific pair of parents creates a “magic” combination.

Sprague and Tatum (1942) proposed the idea of combining ability… general combining ability is the average performance of a strain in a set of cross combinations… whereas specific combining ability is used to describe situations in which specific combinations perform relatively better or worse than would be anticipated based on the average performance of the lines involved. (Kumar, 2024, p. 12).

General Combining Ability (GCA) is a measure of the average performance of a parent line when crossed with a wide variety of other parents (testers). A parent with high GCA reliably produces good offspring, no matter which other parent it’s crossed with. This makes it a valuable parent for broad breeding programs and population improvement.

Specific Combining Ability (SCA), in contrast, measures the performance of a *specific* parental cross. High SCA occurs when two particular parents, when crossed together, produce offspring that are significantly better than expected based on their GCA alone. This is the “lightning in a bottle” that breeders seek for creating elite single-cross hybrids.

Student Note: Remember that GCA is a property of an individual parent, while SCA is a property of a specific *cross* between two parents.

Unpacking the Genetics: Additive vs. Non-Additive Gene Action

GCA and SCA are the outcomes breeders observe; the underlying causes are found in genetics, specifically in the types of gene action that control a trait like yield.

Professor’s Insight: Understanding the type of gene action for a trait (like grain yield or pest resistance) dictates the entire breeding strategy. You can’t just “select for” SCA; you have to find it through systematic test-crossing.

…general combining ability allowed breeders to take advantage of the variability already present… While SCA is used to identify promising single crosses… Additive variance and additive additive interaction variance are the causes of GCA variance. SCA variance, on the other hand, results from dominance variance… and dominance dominance components. (Kumar, 2024, p. 12-13).

The thesis explains that these two concepts are directly linked:

  1. Additive Gene Action is the cause of GCA. In this model, the value of a trait (like height) is the sum of the contributions of individual alleles. Each “good” allele adds a small, positive effect. This type of variation is highly heritable and predictable. A parent with high GCA has accumulated many of these positive, additive alleles and passes them reliably to its offspring.
  2. Non-Additive (Dominance) Gene Action is the cause of SCA. This occurs when the hybrid (heterozygous) state at a gene locus is superior to either of the homozygous parent states. This effect is not additive; it’s an interaction at a specific gene locus. This is the primary driver of heterosis, but because it relies on heterozygosity, it is *not* heritable and “breaks” in the next generation. This is why farmers must buy new hybrid seed every year.

Student Note: For an exam, link these pairs: GCA is caused by Additive gene action and is used for population improvement. SCA is caused by Non-Additive (Dominance) gene action and is used to identify elite F1 hybrids.

ConceptBreeding TermGenetic CausePrimary Use
Predictable / HeritableGCA (General Combining Ability)Additive Gene ActionSelecting good parents
Unpredictable / Non-HeritableSCA (Specific Combining Ability)Non-Additive (Dominance) ActionIdentifying elite hybrid crosses

Table 1: The relationship between breeding concepts and their underlying genetic causes.

The Goal of Hybrid Breeding: Understanding Heterosis

The entire purpose of studying combining ability is to find and maximize heterosis. Heterosis, or “hybrid vigor,” is the phenomenon where the hybrid offspring (F1 generation) outperforms both of its inbred parents in traits like yield, size, or stress tolerance.

Professor’s Insight: Heterosis is the economic engine of the modern seed industry. The discovery of how to exploit it in maize in the early 20th century revolutionized agriculture.

Shull (1908) first presented the idea of heterosis… Since 1960, significant progress has been achieved in the testing and development of inbred lines for the creation of hybrids, which has significantly boosted the yield advantage of maize (Budak et al. 2002). … The findings of numerous studies… supported the idea that heterosis is caused by the accumulation of many favorable dominant genes… (Kumar, 2024, p. 20-21).

The thesis discusses the two main genetic hypotheses for heterosis (p. 21):

  1. The Dominance Hypothesis: This is the most widely accepted view. It suggests that inbred parents accumulate harmful, recessive alleles. When two different inbreds are crossed, each parent provides dominant, favorable alleles that mask the harmful, recessive alleles from the other parent. The result is a hybrid that is fitter than both parents. This aligns perfectly with the concept of SCA.
  2. The Over-dominance Hypothesis: This hypothesis suggests that the heterozygous state (e.g., ‘Aa’) is intrinsically superior to either homozygous state (‘AA’ or ‘aa’).

In practice, breeders measure heterosis in three ways. The most commercially important is Standard Heterosis, which compares the new hybrid’s yield not to its parents, but to the best commercial hybrid (the “standard check”) already on the market (p. 65).

Student Note: Remember the three types of heterosis: Average heterosis (superior to the parents’ average), Heterobeltiosis (superior to the *better* parent), and Standard heterosis (superior to the *commercial check*).

The Real-World Challenge: Genotype x Environment (GxE) Interaction

A breeder’s work is not done once they find a high-SCA cross. That hybrid must perform well everywhere a farmer might plant it. This is the challenge of Genotype x Environment (GxE) Interaction.

Professor’s Insight: GxE is arguably the biggest problem in applied breeding. A hybrid that is #1 in a high-rainfall, high-fertilizer research station might be #10 in a drought-prone, low-input field. Stability across environments is just as important as peak yield.

Genotype x Environment interactions present a significant challenge in cultivar development and selection for a given region or location. …relative rankings of genotypes frequently change when compared across different regions or settings. (Kumar, 2024, p. 3).

This thesis, like most breeding studies, evaluates hybrids across multiple environments (different locations and seasons) to test their *stability*. A hybrid’s performance is not a single number; it’s a response to different conditions. Breeders use statistical models, like the one developed by Eberhart and Russell (1966), to quantify this stability (p. 25).

A “stable” genotype is not one that yields the same everywhere. It is one that performs predictably. An ideal, stable hybrid has a high average yield, a regression coefficient (bi) of 1.0 (meaning it scales perfectly with the environment’s quality), and a deviation from regression (S2d) of 0 (meaning it never has an unexpected bad performance). Farmers depend on this stability for their livelihood.

Student Note: A genotype is considered stable if it has a high mean yield, a regression coefficient (bi) near 1.0, and a deviation from regression (S2d) near zero.

This content has been reviewed and edited by the Professor of Zoology editorial team. All explanatory text, excluding direct quotations from the source thesis, is original content developed for educational use.


Key Takeaways

  • Combining Ability is Predictive: Breeders use GCA (General Combining Ability) to find good average parents and SCA (Specific Combining Ability) to find elite specific hybrid crosses.
  • Genetics Defines Strategy: GCA is driven by predictable, heritable additive gene action. SCA is driven by non-heritable non-additive (dominance) gene action, which creates heterosis.
  • Heterosis is the Goal: Hybrid vigor (heterosis) is the expression of high SCA, most likely caused by dominant alleles from one parent masking recessive harmful alleles from the other.
  • Stability is Essential: Genotype x Environment (GxE) interaction means a hybrid’s performance can change across locations. Breeders must test for stability to ensure predictable yield for farmers.

MCQs (Multiple Choice Questions)

  1. A breeder crosses Parent A with 10 different testers and finds all the offspring are high-performing. Parent A is said to have high…
    • A) Specific Combining Ability (SCA)B) General Combining Ability (GCA)C) HeterobeltiosisD) GxE Interaction
    Correct Answer: B) General Combining Ability (GCA). Explanation: GCA measures the average performance of a parent across many crosses. Since Parent A performed well with all 10 testers, it has a high average value.
  2. Non-additive gene action (dominance) is the primary genetic cause of…
    • A) GCA
    • B) SCA
    • C) Heritable variation
    • D) A stable phenotype
    Correct Answer: B) SCA. Explanation: Specific Combining Ability (SCA) and heterosis are the result of non-additive effects, where the heterozygous combination of alleles is superior to the homozygous parents.
  3. A new hybrid, “Hybrid X,” is compared to “900M Gold,” the best hybrid currently sold. Hybrid X yields 10% more. This 10% increase is called…
    • A) Average heterosis
    • B) Standard heterosis
    • C) Regression coefficient
    • D) Additive variance
    Correct Answer: B) Standard heterosis. Explanation: Standard heterosis (or economic heterosis) compares a new hybrid’s performance to a commercial “standard check,” which is the most relevant measure for farmers.

Frequently Asked Questions (FAQs)

What is a ‘tester’ in maize breeding?
A tester is a known inbred line or population used as a common parent in a series of crosses (a “line x tester” design) to evaluate the combining ability of new, unproven lines.

Why can’t farmers just replant seeds from a hybrid?
Because heterosis is caused by non-additive gene action (SCA), the F1 hybrid’s genes segregate in the F2 generation. This “breaks” the elite combination, resulting in low and variable yields.

What is additive gene action?
It’s when the total effect of genes on a trait is the simple sum of their individual effects. This is predictable and can be “fixed” in a population through selection over time.

What is the Eberhart and Russell (1966) model used for?
It is a statistical model used to analyze Genotype x Environment (GxE) interaction and determine the *stability* of a hybrid across multiple locations and years.

Lab / Practical Note

To perform a “line x tester” cross in the field, you must control pollination. This involves covering the female ear shoots of one parent (the “line”) with a shoot bag *before* its silks emerge to prevent contamination. Later, pollen is collected from the male tassel of the other parent (the “tester”) in a tassel bag. This pollen is then carefully applied to the silks of the “line,” and the ear is re-bagged and labeled. This process is the foundation of all hybrid breeding programs.

For further reading on the methods and concepts discussed in this thesis, explore these high-authority resources:

Primary Source: Kumar, Indrajeet. (2024). GENETIC AND MOLECULAR CHARACTERIZATION OF MAIZE. Doctoral Thesis (PhD in Zoology), Maharaja Agrasen Himalayan Garhwal University, Uttarakhand, India. Supervised by Dr. Sachin Chaudhary & Dr. Satyandra Kumar. Pages used: 1-199.

Note: The provided PDF document includes several published articles (e.g., Pages 20-34) embedded within the thesis structure. All in-text citations (Kumar, 2024, p. X) refer to the thesis pagination provided in the PDF file.

If you are the original thesis author and wish to provide corrections or context, please contact us at contact@professorofzoology.com. We invite universities and researchers to contact us for opportunities to host and promote official thesis abstracts and research summaries.


Indrajeet Kumar, PhD (Zoology), Maharaja Agrasen Himalayan Garhwal University.

Reviewer: Abubakar siddiq, PhD, Zoology.

Note: This summary was assisted by AI and verified by a human editor.

Disclaimer: This article is an academic summary for students and researchers. It is not the original thesis but an educational interpretation of its core concepts.



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