THE PLANT FAMILY SOLANACEAE:

FRUITS IN SOLANACEAE (mostly after Hunziker Ref. 569a)

(J.A.M.van BALKEN)






Fruits originate from the female reproductive part of a flower the gynoecium and are ripenened and seed bearing ovaries (ovary). In the plant family Solanaceae the gynoecium is in most cases bicarpellate ( two carpels) with the following few exceptions:
1) Unicarpellate: in the genus Melananthus Walp. (1850) most probably?
2) 2 (3-4) Carpellate: in the genus Capsicum L.
3) 3-5 Carpellate: in Nicandra Adans (1763), in two species of Jaborosa Juss., and in Trianaea Planch. et Linden.
4) 4 Carpellate : in one species of Iochroma Benth. (1845) (Iochroma umbellatum (Ruiz. et Pavon.) Hunz.).
The number of locules in the ovary are normaly in accordance with the number of carpels (exceptions: Datura L. 2-carpellate and 4 locular due to false septa, some species in the Tribe Lycieae Hunz. such as in Grabowskia Schltdl. in its upper part, and Vassobia Rusby in part of the lower half.
As well as there is a relation between the number of carpels and the number of locules, there is also a relation between the number of ovules in the ovary and the number of seeds in the fruits. The ovule-orientation in Solanaceae is anatropous (Metternichia Mik., Nierembergia Ruiz. et Pav., etc. , anacampylotropous (Phrodos Miers, Grabowskia Schltdl., Vassobia Rusby), hemitropous (Cestrum L. etc.) or hemicampylotropous (Capsicum L., Schizanthus Ruiz. et Pavon, Lycium L.etc.), being born on a rather fleshy placenta, usually many in each locule, sometimes few (in Grabowskia Schltdl. two pairs in each carpel, or one pair in some Lycium L. species), by exception a solitary ovule in Melananthus Walp., generally with similar size (exception: Grabowskia Schltdl. with two unequal elongated ovules in each locule, the upper one larger than the lower one).

The fruits in Solanaceae are in most cases berries or capsules (including pyxidia) and to a lesser extend drupes.

Berries
1) Subfamily CESTROIDEAE Schltdl.(only Cestrum L. and Latua Phil.)
2) Subfamily JUANULLOIDEAE (Hunz.) Hunz.(except Markea Rich. and Rahowardiana D'Arcy)
3) Subfamily SOLANOIDEAE Schltd. except:
    a. Datura L. species (but one species havong a berry: Datura ceratocaula Ort.),
    b. Oryctus S. Watson (capsule)
    c. two species of Lycium L.(drupe)
    d. Grabowskia Schltdl.(drupe),
    e. tribe Hysocyameae Endl.(pyxidium)
4) Subfamily ANTHOCERCIDOIDEAE (G. Don) Tétényi (only the genus Duboisia R. Br.).

Capsules
1) Subfamily CESTROIDEAE Schltdl. (except Cestrum L., Latua Phil.)
2) Subfamily JUANULLOIDEAE (Hunz.) Hunz. (only Markea Rich. and Rahowardiana D'Arcy)
3) Subfamily SOLANOIDEAE Schltd. (only Datura L.[except D. ceratocaula Ort. ] and Oryctus S. Watson)
4) Subfamily SALPIGLOSSOIDEAE (Benth.) Hunz.
5) Subfamily SCHIZANTHOIDEAE (Miers) Hunz.
6) Subfamily ANTHOCERCIDOIDEAE (G. Don) Tétényi (except Duboisia R. Br.)
7) Tribe Hysocyameae Endl.(Pyxidium)

Drupes
1) Subfamily SOLANOIDEAE Schltd.,
    a. tribe Lycieae Hunz. (two species of Lycium L.; Grabowskia Schltdl.)
    b. tribe Solaneae Miers, subtribe Iochrominae (Miers) Hunz.


 

BERRIES

FORM

The form of the berries is globose (nearly sphaerical; e.g. Solanum pseudocapsicum L. ), sphaerical, globose-depressed (to more or lesser extend; e.g. Atropa belladonna L.), subglobose( e.g.Solanum luteum Miller), subsphaerical, subglobose-depressed, ovoid, ovate-globose, ovoid-oblong, elongated, ellipsoid (e.g.Solanum trachycarpum Bitter ex. Sodori), obovate, pyriform (pear-shaped; e.g. Lycopersicon esculentum Miller.var. pyriforme), narrowly pyriform, fusiform (spindle shaped; e.g. Brugmansia versicolor Lagerh.), umbilicate (fruits with a small central depression), or conical (cone-shaped), The berries are most of the time pleuriseminate (many-seeded, with up to 1600-1800 seeds) such as the fruits of some Solanum species) or pauciseminate (few-seeded,1-10 seeds), such as in some Cestrum species, Dyssochroma, Nothocestrum, some Physalis species, Nextouxia and Duboisia

Lycopersicon esculentum Miller var pyriforme (Pyriforme) Solanum trachycarpum Bitter ex. Sodori (ellipsoid) Solanum luteum Miller.(almost globose)
Atropa belladonna L.(globose-depressed) Solanum pseudocapsicum L. (globose)

Brugmansia versicolor (fusiform)

Courtousy of the American Brugmansia and Datura Society

 


 

PLACENTATION

This term refers to the arrangement of ovules in the ovary; indicates the pattern of carpellary fusion. In Solanaceae this is mostly axial (axial placentation).. The number of locules are normally in accordance with the number of carpels and mostly bi-locular in Solanaceae. Exeption is made for the pluricarpellate gynoecia in cultigens (e.g. Lycopersicon esculentum Miller cultivar). As well as there is a relation between the number of carpels and the number of locules, there is also a relation between the number of ovules in the ovary and the number of seeds in the fruits. The placenta is mostly axial (Cyphomandra betacea (Cav.) Sendt.), though there are exeptions (basal/apical e.g.Capsicum annuum L.).

Lycopersicon esculentum Miller cultivar (pleuricarpellate, tetra-locular) Cyphomandra betacea (Cav.) Sendt, Berry ellipsoid. Pleuriseminate. Bilocular. Placenta axile
A: Seed; B: Locule (Bilocular); C: Placenta (axial); D: septum; E: Mesocarp + F: Exocarp= Pericarp

Capsicum annum L. Basal placentation

 


 

PERICARP

The pericarp is the "shell"of the fruit ( see diagram) covering the locules or in case of a drupe covering the pit (stone, pyrene). It consists of three layers: the outerlayer called exocarp (ectocarp), the often juicy midlayer or mesocarp and the innerlayer or the endocarp.The pericarp may be juicy and thick, containing sclerosomes (Witheringia L'Heritier, Acnistus Schott, a few Physalis L. species etc., ) or lacking them (the majority of Physalis L. , Jaltomata Schltdl., Lycopersicon Miller), or may be thin without sclerosomes (Chamaesaracha (A. Gray) Benth. et Hook, Quincula Raf.), or thin with small sclerosomes (Darcyanthus Hunz.), or, by exception, frail, easily breaking into irregular pieces (Quincula Raf., Chamaesaracha (A. Gray) Benth. et Hook). The Exodeconus Raf.berries have, in two species, their apical sector sclerified up to the middle, but pericarp sclerification attains its highest development in the drupaceous fruits of Grabowskia Schltdl., Lycium cooperi A. Gray, Lycium ameghinoi Speg., etc.; in these cases the mesocarp remains fleshy, but the endocarp sclerifies completely, giving rise to two pyrenes enclosing one or various seeds. Finally the fruit of Lycianthes lycioides (L.) Hassler has up to eight 1 or 2 seeded pyrenes.


 

SCLEROSOMES

Scleroids (stone cells) are groups of cells in the pericarp whose walls are thickened with lignin. Some Solanaceae species contain many some few- or lack scleroids.

Solanaceae species lacking scleroids:

Cuatresia Hunz., Jaltomata Schltdl., most Physalis L. species, Quincula Raf, Chamaesaracha (A. Gray) Benth. et Hook, Deprea orinocensis (Kunth) Raf., Exodeconus integrifolius (Phil.) Axelius, Tubocapsium (Wettst.) Makino, most Capsium L.species, Athenea Sendtn., Solanum L.(sections Pseudocapsicum (Moench) Roem. et.Schult., Chamaesarachidium Bitter, Glaucophyllum Child., Lyciosolanum Bitter , Leptostomonum (Dunal) Bitter), Cyphomandra Sendtn., Lycopersicon Miller and Normania Lowe.

Solanaceae species with only a few scleroids:

Brachistus Miers, Phrodus Miers (2 in distal pole mesocarp), Exodeconus pusillus (2), Physalis L. (Physalis alkekengi L., 1-11 up to 20-30; Physalis. carpenteri Rydb. 1 to various), Capsium L. (up to 5 in Capsicum eximium Hunz., Capsicum hookerianum (Miers) OK), Aureliana Sendtn. (few ca 4 or up to 18), Larnax Miers (few up to30 or none), Dunalia Kunth. (not many, exceptionally none), Withania Pauq. (few 3-6), Solanum L. (section Campanulisolanum Bitter) and Lycianthus (Dunal) Bitter (more freqeunt, but not always 1-10).

Solanaceae species with many scleroids:

Deprea sylvarum (Standl. & CV Morton) Hunz, Leucophysalis Rydberg, Iochroma Benth, Eriolarynx (Hunz.) Hunz., Vassobia Rusby and Solanum L. section Archaeosolanum Marzell.(up to 70);


 

CAPSULES

Capsules are mostly dehiscent dry fruits composed of two or more carpels, generally with several or many seeds in each carpel. The capsule dehiscence may be septifrague with two or four valves (e.g.Datura strammonium L.), or usually septicide [(septicidal), dehiscence is along the septa, the area where the carpels or locules are fused to each other, separating the carpels (e.g. Petunia Jussieu , Fabiana Ruiz et. Pav.], loculicide [(loculicidal) the capsule dehiscence is via slits formed in the outer wall of the locules (or cavities)],or septicide-loculicide [(septicidal-loculicidal) Salpiglossoideae (Benth.) Hunz, Metternichia Mik., Nicotiana L. , Anthocercidoideae (G. Don ) Tétényi )]. In Tribe Hysocyameae Endl.the capsule (Pyxidium or Pyxis) opens circumscissle .

Caspsule dehiscence:

Loculicidal, septicidal and circumscissile

Datura strammonium L. Dehiscence septrifrague.

Courtesy: Robert H. Poppenga, DVM, PhD

 


 

DRUPE

Drupes are fleshy fruits, generally grey, brown, or another dull color, with a very thin and dry ovary wall, so that the food is largely confined to the seeds. These may be further subdivided based on the number of seeds and whether the fruit remains closed at maturity ( indehiscent ) or opens naturally ( dehiscent ). There are only a few examples in the plant family Solanaceae:

1) Subfamily SOLANOIDEAE Schltd.,
    a. tribe Lycieae Hunz. (two species of Lycium L.; Grabowskia Schltdl.)
    b. tribe Solaneae Miers, subtribe Iochrominae (Miers) Hunz.

Diagram of a drupe.

 


 

 

CALYX ACCRESCENCE

When the calyx is accrescent ,this one being a common feature with different var. degrees of presence in the family Solanaceae, it may be rigid (Hyoscyamus niger L.)or not (Atropanthe Pascher), smooth (Scopolia Jacq.) or ribbed (Anisodus Link), inflated with its upper sector almost closed ( Physalis mendocina Phil. , Quincula Raf.)or tightly appresed against the pericarp (Darcyanthus Hunz.), or loosely enclosing it with the upper sector open (Chamaesaracha (A.Gray) Benth. et Hook., Leucophysalis Rydberg), or rotate to reflexed never enclosing the berry (Jaltomata Schltdl., etc.).

Hyoscyamus niger L.Fruit a pyxidium. Calyx accrescent and rigid. Physalis mendocina Phil..Calyx accrescent and completely envelloping the fruit.

 

 

Accrescent calyces:

Enveloping, enclosed by or surrounding the berry in variable ways.

Latua Phil., Browallia L., Juanulloideae (Hunz.) Hunz, Hawkesiophyton Hunz, Archeaosolanaum Marzell, Lycopersicon Miller, Juanulloa Ruiz. et Pavon, Merinthopodium J. Donn. Sm., Rahowardiana D'Arcy, Schultesianthus Hunz., Trianaea Planch. et Linden , Schwenckieae Hunz.(Protoschwenckia Soler., Heteranthia Nees et. Mart), Brachistus Miers , Deprea Raf. (loosely enveloping and inflated), Grabowskia Schltd.(partially), Exodeconus Raf.(inervation much conspicuous), Athenaea Sendtn. (fruiting calyx up to 7x the floriferous ones), Darcycanthus Hunz.(except its upper surface), Eriolarnyx (Hunz.) Hunz,(in part or completely), Nicandra Adans, Phrodus Miers, Hyoscyamus L. (completely hidden in urceolate or obconic calyx), Anisodus Link.(enclosing tightly, calyx ribbed), Atropanthe Pasher (pericarp hidden in enlarged, open, urceolate, bladdery calyx), Physochlaina G. Don (pericarp hidden in much accrescent inflated calyx), Przewalskia Maxim (pericarp contained in the spacious, much accrescent calyx), Scopolia Jacq., (enclosed rather closely in the not bladdery, slightly longer calyx), Salpiglossis Ruiz. et Pavon (surpassing the berry or slightly shorter), Reyesia Gay, Mandragora L. (shorter or as long as), Larnax Miers (completely by the cardiform, urceolate calyx), Triquera Cav., Normania Lowe (enclosed by a calyx with leafy lobes), Jaborosa Juss.(partly enclosed), Jaltomata Schltdl.(not enclosing; calyx rotate and reflexed), Nothocestrum A. Gray (variable), Chamaesaracha (A.Gray) Benth. et Hook.(thightly invested) and Oryctes S. Watson (closely fitting the berry).

Slightly accrescent calyces:
Leucophysalis Rydberg (calyx thin campanulate or slightly urceolate, tightly appressed to the berry or loosly enclosing it, not angled or ribbed, always open in the upper sector), Iochroma Benth.( usually addressed to the basal half of the pericarp or to its entirety, irregular cut in lobs or segments due to the growing berry), Vassobia Rusby, Saracha Ruiz. et Pavon, Cyphomandra Sendtn., Atropa L. (fruiting calyx somewhat enlarged, its lobes clearly spreading) and Salpichroa Miers.

Not accrescent calyces:
Datura L.(persitent in a collar), Cuatresia Hunz., Discopodium Hochst., Dunalia Kunth, Acnistus Schott. (calyx not accrescent or enlarging significantly), Tubocapsicum (Wettst.) Makino, Capsicum L., Aureliana Sendtn.and Anthocercidoideae (G. Don ) Tétényi




GLOSSARY ( partly Ref. 946a)

Accrescence: Enlarging of the calyx during maturation of the fruits.

Anatropous: Ovule orientation. An ovule that is inverted and straight with the micropyle facing the placenta

Anacampylotropous: Ovule orientation somewhere in between anatropous and campylotropous.

Axial placentation: The ovules (seeds) are locate along the axis

Basal (apical) placentation: The ovules (seeds) are located at the base or at the apex of the ovary or fruit

Berries: One or more carpels developed within a thin covering, very fleshy within, with the seeds embedded in the common flesh of a single ovary, such as in Lycopersicon esculentum (tomato).

Calyptra : Hood or cap-like part covering the calyx of certain plants.

Campylotropous: Ovule orientation. Ovule oriented transversely, i.e. with its axis at right angles to its stalk, and with a curved embryo sac.

Campanulate: Bell-shaped; cup-shaped with a broad base.

Capsule: Dehiscent fruit composed of two or more carpels, generally with several or many seeds in each carpel.

Cardiform:Calyx is in the form of a heart.

Carpels: Carpels are structures that are made up from an ovary and a stigma and that contain one or more ovules.

Chalaza: Part of an ovule to which the end of the stalk (fungible) is attached.

Circumscissle: Capsule or pyxis that dehisces circumferentially. Opening or divided by a horizontal circular line; applied to pods or seed capsules that open with a lid, like a box. The fruit in such cases is called pyxidium.

Conical:Cone-shaped; widest at the base and tapering to the apex.

Cuneiform: Cuneatus or wedge-shaped: with straight sides converging at the base. (late cuneatus when they make an angel of 90 degrees or more and anguste cuneatus when they make an angle of less than 90 degrees)

Dehiscence: The fruit opens naturally (opposite remains closed at maturity ( indehiscent )

Drupes: ("stone fruit") A simple fruit produced from a single carpel, usually one-seeded, with an outer fleshy layer of tissue called the pericarp and an inner, heavy stony layer called the endocarp .

Dry fruits: Generally grey, brown, or another dull color, with a very thin and dry ovary wall, so that the food is largely confined to the seeds. These may be further subdivided based on the number of seeds and whether the fruit remains closed at maturity ( indehiscent ) or opens naturally ( dehiscent ).

Ectocarp: (see exocarp) Outmost layer of pericarp

Ellipticus: (ovalis,oval): Elliptical, acute at each end.SADT numbers 3-4; lenght: breadth 2:1 to 3:2 with sides equally curved from middle.

Ellipsoideus:Ellipsoidal: a solid with a elliptical figure.

Elongated:Elongatus; Long, longer.

Endocarp: Innermost differentiated layer of pericarp.

Exocarp: (or ectocarp); Outermost layer of pericarp.

Fleshy fruits: are usually juicy and brightly colored, contrasting with their background to make them more noticeable to animals, who are responsible for their dispersal. All fleshy fruits are indehiscent and considerable fleshy tissue is developed as the ovary changes into the fruit.

Fruit form: The form (look ) of a fruit as pictured in the diagram below.

A: Cuneiform; B:Fuseiform; C: Ellipticus (ellipsoideus); D: Pyriformis; E: Obpyriformis; F: Obovatus (obovoideus); G: Ovatus (ovoideus); H: Ovalis: I: Sphaericus (globosus); J: Subglobosus; K: Globose-depressed.

 

Fruits: Are the ripened and seed-bearing ovaries of flowers. Fruits are nearly as varied in color, form, size, texture, and number as are flowers, making them valuable tools in plant identification. One can distinguish dry fruits (e.g. capsules) and fleshy fruits (such as beriies and drupes).

Funicle: Seed stalk or ovule stalk

Fuseiform: Fusiform, swollen at middle and tapering to each end like a spindle , narrowly ellipsoid.

Globose-depressed:Round like a ball, sphaerical but one end a little bit flattened.

Globosus:Round like a ball, sphaerical.

Gynoecium: The female reproductive organs, the gynoecium, include the carpels. Carpels are structures that are made up from an ovary and a stigma and that contain one or more ovules. One (unicarpellate) or more carpels (bi, tri, etc. carpellate) may be combined to a pistil (ovary, style, stigma), the gynoecium as a whole.

Hemitropous: Half-inverted between anatropous and campylotropous.

Hemicampylotropous: Ovule orientation. Like campylotropous but having the raphne terminating half way between the chalaza and the orifice.

Inflated:: Calyx bladdery, i.e. thin, membranous and swollen.

Integument: Covering layer; internal integument and external integument.

Locules: Compartment or cavity of an ovary, anther, or fruit.

Loculicidal: Capsule that dehisces longitudinally into the cavity of the locule

Mericarp: A portion of fruit that seemingly matured as a separate fruit.

Mesocarp: Middle layer of pericarp.

Micropyle: Opening or mouth (orifice)of a ovule.

Nucleus: Nucleus the body that contains the genetic material of the egg/ovule..

Obconic: Conical with the apex downwards.

Obovatus: Reversed ovate, the broadest part above the middle , lenght to breadth about 3:2.

Obovoideus: As ovoidal but broadest part above the middle..

Obpyriformis: As pyriform but the broadest part above the middle.

Orifice:Opening; mouth of the ovule.

Ovalis: Oval, elliptic, i.e. the broadest at the middle, the sides curved and lenght : breadth usually 2:1.

Ovary:: The part of the pistil that contains the ovules.

Ovatus: Ovate, oblong or elliptical, broadest at the lower end , so as to resemble the longitudinal section of an egg. SADT nos. 38-39; lenghth : breadth = 2:1 to 3:2, broadest below the middle.

Ovate-oblong: A form in bewteen ovatus and oblong, oblong being elliptical, obtuse at each end. SADF nos. 15-15; lenght : beadth = 2:1 to 3:2, the sides almost parallel.

Ovoideus:(Ovoidal) A solid with an ovate figure, or resembling an egg.

Ovoid-oblong: A form in beteen ovoides and oblong.

Ovule orientation: The way the ovules are arranged on the placenta.

R: Raphe: The part of the stalk of an anatropous ovule that is fused along the side of the ovule.
M: Micropyle; Opening or mouth (orifice)
F: Funicle (stalk);
N: Nucleus;
C: Chalaza; part of an ovule to which the end of the stalk (fungible) is attached.
II: Integument internal;
IE: Integument external.

Ovules: The body which, after fertilization, becomes the seed.

Pauciseminate: Few-seeded.

Pericarp: Fruit wall consiting of three layers: exocarp, mesocarp and endocarp..

Pistil: The seed-bearing organ of the flower, consisting of the ovary, stigma, and style when present

Pit: (stone; pyrene).The stone of a succulent fruit made up of the endocarp and the seed.

Placenta: Region of attachment of seeds on inner fruit wall.

Pleuriseminate: Many-seeded.

Poricidal: Capsule that dehisces through pores

Pyriformis:Pear-shaped.

Pyxidium: A pod which divides circularly into an upper and lower half, of which the former acts as a kind of lid (operculum).

Raphe:The part of the funiculus that is fused along the side of the ovule.

Reflexed (Reflexus) Bent abruptly backwards at more than 90 degrees.

Replum:. Persistent septum after dehiscence of fruits.

Rostellum: ( Beak). Persistent stylar base on fruit.

Rotate:Describes a calyx or corolla that is wheel-shaped flat and circular in outline.

Scleroids: (stone cells) Scleroids are groups of cells whose walls are thickened with lignin.A variably shaped sclerenchymatous cell of a higher plant.

Seeds: Matured ovules, consist of an outer coat or wall, which is usually very tough, hard, or woody, within which are cotyledons and the embryo. Seeds normally have just one embryo, but sometimes have more than one, which results in two or more new plants growing from one seed. Seeds are developed as a result of the fertilization of the egg in the ovule of the ovary of a flower. Typically seeds are oval or globular and range in size. Seeds vary greatly in color, texture, longevity, and methods of dispersal. Some of the modifications of seeds which aid in dispersal are coverings of spines, hooks, bristles, cotton, or plumes, or having wings and arils. They also vary in the types and abundance of food they contain.

Septicidal: Capsule that dehisces longitudinally through the septa

Septicidal-loculicidal: Dehiscence partly septicidal, partly loculicidal.

Septifragal: (or valvular) Capsule, one with valves breaking away from the septa,

Septum: (Dissepiment). Partition. For example in a fruit partition of the fruit in locules.

Sphaericus: (Globose) Forming nearly a true sphere.

Subglobosus:. Sub meaning almost. Almost globose.

Subglobose-depressed: Almost globose depressed.

Subsphaerical: Almost sphaerical.

Umbilicate: Navel-like, having a small central depression or hollow.

Urceolate:Hollow and cylindrical or ovoid, and contracted at or below the mouth, like an urn.

Valvular: (Septifragal) Capsule, one with valves breaking away from the septa,


 

 

 

 

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