berry [ BER-ee ] noun, plural berries: a fleshy, simple fruit without a stone or core
hesperidium [ hes-puh–RID-ee-uhm ] noun: a berry with a leathery rind and segmented flesh such as citrus fruit
pepo [ PEE-poh ] noun: an epigynous berry with a hard, inseparable rind common to the gourd family
Photo © Christa Watters
Can you identify any berries among the edibles in these pictures? Citrus fruits are berries called hesperidiums. Although nutritionists consider eggplants, peppers, pumpkins, squash, and tomatoes vegetables, botanists categorize them as fruits and they too are berries. Melons, pumpkins, and squash are berries called pepos. Other edible berries include avocados, bananas, blueberries, cucumbers, grapes, kiwis, maypops, pawpaws, and persimmons.
A berry is a fleshy fruit that develops from a single ovary of a flower. The ovary wall develops into the pericarp that protects the seeds inside. The pericarp has three layers: exocarp (outermost layer), mesocarp (middle layer), and endocarp (innermost layer). Where the ovary is located in relationship to the other flower whorls (androecium/stamens, corolla/petals, and calyx/sepals)—whether it is above (superior) or below (inferior) their point of attachment—as well as the substance of the pericarp distinguishes the four berry fruit types. A fruit formed from an inferior ovary may include other flower parts, such as the hypanthium (the floral cup/tube or the enlarged receptacle), petals, sepals, or bracts. Berries do not naturally dehisce.
Superior Ovary
In a hypogynous flower, the (a) androecium, (p) petals, and (s) sepals arise from the (r) receptacle beneath the (g) gynoecium. The ovary is superior.
“True” Berry Cross Section
A “true” berry forms from a superior ovary with one carpel or multiple fused carpels. It has a fleshy pericarp and usually many seeds. Examples include: avocado (with one large seed), eggplant, grape, kiwi, pawpaw, pepper (air replaces the flesh), persimmon, and tomato (pictured above).
Hesperidium Cross Section
A hesperidium is a modified berry of the genus Citrus. A leathery exocarp, commonly called a rind, peel, or zest, encloses the pulp, which is comprised of 8–16 carpels/locules with fluid-filled vesicles partitioned by septa. Examples include grapefruit, lemon (pictured above), and orange.
Inferior Ovary
In an epigynous flower, the androecium, petals, and sepals are attached to the top of the ovary, which is inferior.
Epigynous Berry Cross Section
An epigynous berry (sometimes called a “false” berry) is an accessory fruit formed from an inferior ovary and other flower parts. Examples include banana, blueberry (pictured above) and cranberry.
Pepo Cross Section
A pepo is an epigynous berry common in the Cucurbitaceae, the gourd family, formed from an inferior ovary and with a hard or firm, inseparable rind. Examples include cantaloupe, cucumber, pumpkin, squash, and watermelon (pictured above).
Other EDIBLE Berries
Left to right: Asimina triloba , Cucurbita moschata, Diospyros virginiana, Poncirus trifoliata, Passiflora incarnata.
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Some POISONOUS Berries
NOTE: Plants in the wild should not be eaten without consulting an expert or authoritative field guides for information on identification and food preparation. It is easy to confuse plants in the wild, so you should be 100% sure they are edible before consuming them.
References
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Erwin J. Plant Grower. Blueberry (Vaccinium corymbosum). Department of Plant Science and Landscape Architecture, University of Maryland. (accessed February 3, 2024).
Gandolfo MA, Paolillo DJ, Svitko JL. Topic 14: Flowers, Fruits, and Seeds. Plant Anatomy Online Lab Manual. © 2019 CU-PAC Cornell University. (accessed February 24, 2024).
Mann LK. 1943. Fruit Shape of Watermelon as Affected by Placement of Pollen on Stigma. Botanical Gazette, 105(2): 257–262.
Matheyambath AC, Padmanabhan P, Paliyath G. 2016. Fruit Morphology. Citrus Fruits. Encyclopedia of Food and Health. pp. 136–140.
Nickrent DL. Updated October 2022. Lecture 24 Fruits. Plant Anatomy. Southern Illinois University.
Pao S, Fellers PJ. 2003. Anatomy and Varieties. Citrus Fruits|Oranges. Encyclopedia of Food Sciences and Nutrition (Second Edition). pp. 1341-1346.
Spjut RW. 2003–2015. A Systematic Treatment of Fruit Types. ©The World Botanical Associates.
Stover-Blackburn J, Editor. Fruit Anatomy. UCDavis Fruit & Nut Research & Information Center. © 2024 Regents of the University of California (accessed February 14, 2024).
Weakley AS, Ludwig JC, Townsend JF. 2012. Flora of Virginia. Botanical Research Institute of Texas. ISBN 10: 1889878383. ISBN 13: 9781889878386.
Witcher CL. 2011. Blueberry Fruit Development and Splitting. The Aquila Digital Community. The University of Southern Mississippi.