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Solanum pimpinellifolium

Solanum pimpinellifolium, commonly known as the currant tomato, is a wild relative of the cultivated tomato (S. lycopersicum) in the family , characterized as a diploid (2n=24) herbaceous with a sprawling or procumbent habit, pinnatifid non-prickly leaves, yellow flowers, and small, edible red fruits measuring 0.5–1 cm in . Native to western , particularly the coastal regions of and , it thrives in arid, sandy habitats near water sources or field edges from to over 1,000 m elevation, though populations are declining due to habitat loss from , , and . Taxonomically, S. pimpinellifolium belongs to Solanum section , a group of species with brightly colored fruits, and was formerly classified under the genus with the synonym L. pimpinellifolium (L.) Mill. Its center of genetic diversity lies in northern , where it exhibits high variability in traits such as floral morphology and outcrossing rates, with a self-compatible mating system that includes facultative promoted by features like exserted stigmas and larger flowers. The species has adventive populations on the and extends to northern in wild and feral forms, but it is not truly wild in many areas and can behave as an invasive weed in disturbed sites. As a key genetic resource, S. pimpinellifolium is widely used in tomato breeding programs to introgress traits for resistance, including to , , late blight (via the Ph-3 gene), and Tomato Yellow Leaf Curl Virus (TYLCV, with a major QTL on ), as well as for insect resistance, tolerance to and , and improvements in fruit quality such as higher (sugar content) and enhanced flavor. Its fruits are occasionally consumed raw, cooked, or dried in native regions and are popular among home gardeners, with commercial seeds available, though the green parts of the plant contain toxic . Conservation efforts are critical, as has led to the loss of up to 23 populations in seven Peruvian valleys, underscoring the need to preserve its diversity for future agricultural applications.

Taxonomy

Classification

Solanum pimpinellifolium is classified in the kingdom Plantae, phylum , class , order , family , genus , and species S. pimpinellifolium. It resides within section Lycopersicon of subgenus Potatoe, sharing a close phylogenetic relationship with the cultivated tomato Solanum lycopersicum. This placement reflects its position among the "tomato clade" of wild relatives that form a monophyletic group within the genus. Historically, S. pimpinellifolium was segregated into the distinct genus Lycopersicon as L. pimpinellifolium, a classification established in the 18th and 19th centuries based on morphological distinctions such as fruit characteristics. However, molecular phylogenetic analyses using nuclear genes like granule-bound starch synthase (GBSSI) in the late 1990s and early 2000s revealed that Lycopersicon species are deeply nested within Solanum, prompting their reintegration into the broader genus around 2001–2005. This taxonomic shift was supported by evidence of shared synapomorphies and close genetic affinities among section Lycopersicon members. The binomial authority for Solanum pimpinellifolium is (L.), with the name first validly published in 1755.

Nomenclature

Solanum pimpinellifolium is the accepted binomial name for this wild under the Code of for , fungi, and (ICN), which governs the scientific naming of . The name derives from the Latin solamen, meaning "comfort" or "soothing," reflecting the medicinal uses of some in the , or possibly from sol, referring to the sun-like appearance of certain flowers. The specific epithet pimpinellifolium combines , a in the family (including ), with Latin folium for "leaf," alluding to the resemblance of its leaves to those of Pimpinella . The species was originally described by in his Centuria I Plantarum (1755), where it was established as Solanum pimpinellifolium L. This publication provided the formal , marking its entry into scientific literature. Prior to modern revisions, the species was often classified in the segregate genus , with the basionym (L.) Mill. published by in the eighth edition of The Gardeners Dictionary (1768). Several synonyms reflect historical taxonomic treatments, including Lycopersicon esculentum Mill. subsp. pimpinellifolium (L.) Luckwill and Lycopersicon esculentum Mill. var. racemigerum Hook. f., the latter sometimes treated as a variety distinguished by racemose inflorescences. These names arose during periods when tomatoes were separated from Solanum, but phylogenetic studies since the early 2000s have confirmed its placement within Solanum section Lycopersicon, leading to the current accepted nomenclature. Common names for S. pimpinellifolium include , pimp tomato, and wild cherry tomato, reflecting its small, berry-like fruits and wild origins.

Description

Botanical characteristics

Solanum pimpinellifolium is an annual or short-lived herb with a growth habit that begins erect but becomes procumbent or viny as it matures, reaching heights of up to 3 meters with branches extending similarly. The stems are slender and weak, measuring 8-11 mm in diameter at the base, green in color, and covered with sparse to dense velvety pubescence consisting of trichomes 0.5-1 mm long. It exhibits a simple to branched habit, often sprawling without support. The root system is fibrous, comprising a main root connected to numerous lateral roots that enable to varied conditions. The leaves are interrupted imparipinnate, typically 4-12 cm long and 1.5-8 cm wide, with a hue and velvety pubescence. They feature 5-9 leaflets, including a terminal leaflet that is 2.5-5 cm long and 1-3.5 cm wide with a cordate base, lateral leaflets measuring 1.5-3.5 cm long by 1-2 cm wide, and 1-4 pairs of small interjected leaflets 0.5-1.2 cm long. The pinnate structure and pubescent nature give the foliage a compound appearance similar to that of , reflected in the species epithet "pimpinellifolium." Flowers are small and 5-merous, arranged in racemose inflorescences 4-25 cm long bearing 7-30 blooms. The is stellate, pale to bright yellow, and 1.2-3 cm in diameter, with deeply divided lobes (more than 3/4 of the corolla length) that are narrowly lanceolate and 1-1.5 cm long. The species is self-compatible, facilitating , though can occur due to the floral structure. In comparison to the cultivated Solanum lycopersicum, S. pimpinellifolium has a smaller overall stature, greater branching, and a wild pattern, along with distinct features such as the cordate base of the terminal leaflet and velvety pubescence lacking long trichomes. These morphological traits underscore its status as a wild , contributing to its weedy, sprawling form in natural settings.

Reproduction and fruits

Solanum pimpinellifolium flowers throughout the year in its native subtropical range along the western coast of , where stable environmental conditions support continuous blooming, though the process is influenced by day length, with plants tending to flower later under long-day photoperiods compared to short-day conditions. The hermaphroditic flowers, arranged in scorpioid cymes, are primarily pollinated by native bees capable of , such as bumblebees and other vibration-producing species, which release from poricidal anthers; unlike some , these flowers do not produce and instead offer as the primary reward to pollinators. Successful pollination leads to the development of small, globose berries measuring 0.8-1.2 cm in diameter, which ripen to vibrant red or, in some accessions, yellow hues, each fruit typically containing 10-50 that demonstrate high viability under suitable and conditions. The seeds are minute, 1-2 mm in size, with a reniform (kidney-shaped) morphology, and germinate reliably within 7-14 days when sown at temperatures between 20-30°C, often requiring moist conditions and occasional stratification for optimal emergence. While commonly treated as an in due to sensitivity and seasonal growing cycles, S. pimpinellifolium exhibits variations in the wild, functioning as a short-lived or under favorable tropical or subtropical conditions with minimal disturbance. In natural populations, individual plants can yield up to several hundred fruits, reflecting their adaptation for prolific in coastal habitats.

Distribution and habitat

Geographic range

Solanum pimpinellifolium is native to western , with its primary range encompassing the coastal regions of and southern . This wild species occurs from up to approximately 1,500 m , inhabiting diverse locales including the Andean foothills and arid Pacific coast deserts. In , populations are documented in northern areas like and Lambayeque, as well as southern coastal deserts such as those near and , often along roadsides and field edges. In , it extends from northern tropical forests in Esmeraldas to montane regions in Loja and Azuay, with dense wild stands in undisturbed coastal and inter-Andean valley habitats. The species has a history of early documentation by European explorers in the 18th century, which contributed to its initial botanical descriptions and spread beyond its native range. S. pimpinellifolium has been introduced and naturalized in the Galápagos Islands since the 19th century, where it arrived from continental Ecuador and Peru and now persists as an invasive species in some areas. It is also naturalized in parts of Central America, including Costa Rica, and maintains presence in experimental agricultural plots worldwide for breeding purposes. Wild populations of S. pimpinellifolium span a broad coastal corridor in its native distribution. Despite agricultural expansion, these populations remain relatively widespread along the Pacific seaboard, though fragmented in some northern Peruvian locales.

Ecological preferences

Solanum pimpinellifolium thrives in tropical to subtropical climates, particularly arid to semi-arid conditions prevalent in its native range along the western coast of South America from Ecuador to northern Chile. It prefers environments with annual mean temperatures ranging from 17.9°C to 26.8°C, with a median of approximately 22.7°C, and annual precipitation typically between 1 mm and 2989 mm, though medians indicate low rainfall around 68-70 mm, often with dry months receiving 0 mm. These conditions align with Köppen classifications such as BSk (cold steppe) and BWh (hot desert), supporting its growth in warm, dry microclimates influenced by seasonal fog and occasional high rainfall during El Niño events. The species favors well-drained sandy soils with a range of 3.2 to 8.5, median 7.6, and is adaptable to various soil types including Leptosols and Regosols, excluding heavy Vertisols. It exhibits notable tolerance to saline soils, originating from coastal and brackish habitats, where accessions demonstrate superior root development and retention under high (e.g., EC 31.1 mS cm⁻¹). in preferred soils typically ranges from 0.3 to 1.5 kg/dm³, with a median of 1.4, facilitating root penetration in low-altitude (median 92-93 m) settings. In terms of habitat, S. pimpinellifolium occupies disturbed areas such as dry slopes, plains, riverbanks, and coastal dunes, often associating with other in inter-Andean valleys and coastal regions. Biotically, it interacts with native buzz-pollinating bees, including bumblebees (Bombus spp.) and (Melipona spp.), which vibrate poricidal anthers to release , enhancing set. The shows resistance to certain pests like nematodes and (e.g., at least nine in some accessions) and pathogens, contributing to its persistence in these environments. Key adaptations include through maintained , CO₂ assimilation, and efficient water use under deficit conditions, as seen in accessions like Pim2 and Pim3. It also employs osmotic and ionic adjustments for salt stress, with robust root architecture aiding survival in low-water, saline habitats.

Genetic aspects

Genome structure

Solanum pimpinellifolium is a diploid with a number of 2n=24. Its is approximately 900 in size and consists of 12 chromosomes. The first draft of S. pimpinellifolium was produced in 2012 by the Tomato Genome Consortium as part of the broader sequencing effort, yielding a 739 Mb from Illumina short reads aligned to the 12 chromosomes. A high-quality, chromosome-scale was achieved in 2020 using PacBio long reads and data for the accession LA2093, resulting in an 807.6 Mb with 97.8% completeness and revealing over 92,000 structural variants, including inversions and indels that influence breeding traits. In 2024, a chromosome-level of accession LA1589 was reported, spanning 833 Mb with a contig N50 of 31 Mb and scaffold N50 of 69.7 Mb. Notable genomic features include the presence of key genes such as SP6A, a homolog of the potato StSP6A gene that regulates tuberization and remains functionally intact in S. pimpinellifolium, in contrast to the domesticated tomato where it is disrupted by a stop codon. The genome also harbors loci associated with disease resistance, exemplified by alleles conferring tolerance to Tomato yellow leaf curl virus (TYLCV) introgressed from accessions like UPV16991. Compared to the domesticated tomato S. lycopersicum, the S. pimpinellifolium genome exhibits only 0.6% divergence, reflecting their close relationship and evidence of recent . It shows substantial synteny with the (S. tuberosum) genome, despite over 8% sequence divergence and nine large inversions, underscoring conserved syntenic blocks across the family.

Genetic diversity

Solanum pimpinellifolium exhibits substantial , reflecting its adaptation to diverse coastal environments in western . Studies using high-density (SNP) markers have identified over 24,000 SNPs across 99 accessions from the Tomato Genetics Resource Center (TGRC), highlighting extensive variation within the species. Expected heterozygosity levels range from 0.15 to 0.27 across subpopulations, indicating a high potential for allelic richness, while observed heterozygosity varies from 0.045 to 0.147, influenced by varying degrees of selfing. (SSR) analyses of 248 individuals further confirm moderate to high diversity, with Nei's genetic diversity index averaging 0.40 in northern populations and mean alleles per locus at 8.7. Population structure reveals clinal variation along the Pacific coast, with greater diversity in northern and northern compared to southern regions and the . SSR-based clustering differentiates Peruvian and Ecuadorian populations, with northern Peru showing the highest diversity (Nei's index 0.40) and exclusive alleles, while Galápagos accessions display near-zero diversity due to founder effects and isolation. Genome-wide SNP analyses support this, identifying three single-ancestry and four mixed-ancestry subpopulations, with genetic differentiation correlating to geographic and climatic gradients. Molecular marker studies using SSRs and SNPs demonstrate clear genetic differentiation from the closely related cultivated , Solanum lycopersicum, underscoring S. pimpinellifolium's role as a distinct wild . markers reveal structured variation tied to geography, while data show rapid decay (average 3.3 kb), indicative of historical and recombination. These markers have been instrumental in mapping diversity panels, confirming low between wild and domesticated forms. Genetic variation manifests in key adaptive traits, including size, shape, and resistance, with bioclimatic correlations shaping ecotypic differences. Northern populations exhibit larger s and higher resistance to certain pathogens, while southern ecotypes show adaptations to through traits like reduced stomatal density and enhanced water-use efficiency. For instance, genetic differentiation aligns with and gradients, with southern accessions displaying alleles linked to arid . In conservation genetics, S. pimpinellifolium maintains viability through facultative with rates varying from 0% to 84%, typically up to 40% in natural and higher in northern with exserted stigmas that promote flow. is relatively low, supporting persistence despite self-compatibility, as evidenced by minimal declines in selfed progeny compared to outcrossed ones in related breeding studies. These attributes facilitate effective ex situ preservation in , where hundreds of accessions capture this diversity for future utilization.

Breeding and uses

Role in tomato breeding

Solanum pimpinellifolium, the wild progenitor of the cultivated (Solanum lycopersicum), has been a key resource in tomato breeding programs since the early , with initial interspecific crosses documented as early as the to harness its for crop improvement. As the closest wild relative, it freely hybridizes with cultivated tomatoes, producing vigorous F1 hybrids that facilitate without significant reproductive barriers. This close genetic relatedness has enabled breeders to transfer beneficial alleles from S. pimpinellifolium into elite lines, enhancing resilience in modern varieties. Key traits introgressed from S. pimpinellifolium include to biotic stresses such as tomato yellow leaf curl virus (TYLCV), where monogenic recessive has been identified in accessions like AAU2019, providing a valuable for programs. Similarly, quantitative trait loci (QTLs) for late blight , caused by , have been mapped in accessions like PI 270441, allowing breeders to select for reduced disease severity in cultivated backgrounds. to , induced by , has also been characterized, with combining ability studies showing additive and dominance effects in crosses involving S. pimpinellifolium genotypes. Abiotic stress tolerances, including , are prominent, with S. pimpinellifolium demonstrating superior growth maintenance and yield under compared to cultivated es. It also exhibits , with better and survival under water stress, supporting its use in for arid conditions. Recent studies as of 2024 have identified S. pimpinellifolium regions for improving nitrogen use efficiency in . Breeding methods employing S. pimpinellifolium primarily involve and (MAS) to target regions while minimizing unwanted wild traits. (IL) libraries, such as those developed from accessions like LA2093 or TO-937, have been instrumental in fine-mapping and transferring QTLs for these traits into commercial backgrounds. Notable examples include the hybrid variety 'Mountain Magic', which incorporates S. pimpinellifolium-derived Ph-3 gene for to late blight, resulting in high-yielding, disease-tolerant plants. Despite these advances, challenges persist, particularly linkage drag, where desirable resistance genes are co-inherited with deleterious traits like small fruit size or reduced yield, complicating selection in backcross populations. Innovations using CRISPR/Cas9 have addressed this by precisely editing alleles from S. pimpinellifolium for traits like fruit quality and stress tolerance, reducing linkage drag and accelerating . These tools have targeted multiple genes from accessions to restore ancestral variations beneficial for without broad genomic disruptions.

Culinary and ornamental applications

Solanum pimpinellifolium fruits, commonly known as currant tomatoes, are small (10–15 mm in diameter) and produced in large clusters, making them suitable for fresh consumption in salads, as garnishes on dishes like soups, or in preserves. Their flavor is described as sweet yet tangy with a burst of essence, ideal for adding to salsas and sauces where the sweetness balances spicy or acidic elements. Due to their growth habit and small size, they are not viable for large-scale commercial but are valued in home gardening for direct culinary use. Nutritionally, the fruits are rich in antioxidants, including (up to 39 mg/100 g fresh weight), , and phenolic acids, which contribute to health benefits such as reduced and cardiovascular protection. Vitamin C content ranges from 25 to 50 mg/100 g fresh weight, higher than many cultivated varieties, supporting immune function and synthesis. These compounds are particularly bioavailable when the fruits are cooked, as in sauces or dried forms, enhancing their role in diets focused on disease prevention. In ornamental gardening, S. pimpinellifolium is prized for its vigorous, sprawling vines, attractive foliage, and clusters of brightly colored, pea-sized fruits that add visual appeal to borders or containers. varieties such as 'Red Currant' are popular for their prolific production of vivid red berries, providing both aesthetic value and low-maintenance charm in temperate and tropical gardens. The plant's disease resistance and ease of growth further enhance its suitability as a decorative element without requiring extensive support. Traditionally in the Andean region, indigenous communities have used S. pimpinellifolium for medicinal purposes, including poultices from pulped fruits to treat burns, sunburn, and due to its properties. The high profile also positions it for potential applications in nutraceuticals, such as supplements targeting and oxidative damage. In markets, it occupies a niche in specialty and seed sales, with limited annual production focused on or outlets rather than mass .

Conservation

Status and threats

Solanum pimpinellifolium has not been formally assessed for the of Threatened Species, reflecting its relatively widespread distribution as a wild relative of the cultivated across western . However, populations in its native range are declining overall due to the species' vulnerability in fragmented habitats where human activities predominate, despite its weedy and resilient growth habit allowing persistence in some disturbed environments. The primary threats to S. pimpinellifolium include habitat loss driven by , coastal development, and , particularly in northern and , where industrial farming has overtaken former coastal and arid habitats. Genetic erosion has led to the loss of up to 23 populations in seven Peruvian valleys. exacerbates these pressures by shifting suitable climatic conditions in arid zones, potentially leading to range contraction and increased extinction risk for certain populations. Population trends indicate declines in fragmented areas, with habitat loss contributing to reduced numbers. Legal protections encompass its status as a under the (), with genetic resource access governed by the to ensure equitable benefit-sharing. Ex situ conservation efforts maintain over 1,000 accessions of S. pimpinellifolium in global genebanks, including significant holdings at institutions like the World Vegetable Center and the Tomato Genetics Resource Center, yet these collections are noted to underrepresent the species' full across its native range.

Genetic resource management

Ex situ conservation efforts for Solanum pimpinellifolium primarily involve that maintain living collections for and . The World Vegetable Center (WorldVeg) holds over 320 accessions, representing diverse origins from its native range in coastal and , with ongoing multiplication and evaluation to ensure viability. Similarly, the USDA's Unit (PGRU) in , curates a substantial portion of the species within its broader collection exceeding 6,600 accessions, facilitating global distribution and characterization. techniques, including shoot-tip and encapsulation-dehydration, are employed for long-term storage of wild relatives like S. pimpinellifolium, preserving genetic integrity without recurrent regeneration. In situ and on-farm conservation complement these efforts by protecting natural populations in their coastal habitats. Protected areas in and , such as coastal reserves near and Loja, safeguard wild populations, while collaborative collecting missions with local universities since the have documented and sampled in situ. Community-based programs in these regions promote on-farm maintenance by farmers, integrating wild accessions into traditional agroecosystems to sustain adaptive diversity. Utilization of S. pimpinellifolium genetic resources focuses on pre-breeding to introgress valuable traits into cultivated . Recent development of backcross inbred lines (BILs) from interspecific crosses, as detailed in 2024 studies, has created populations evaluated for heat stress tolerance, serving as bridges for allele mining. These efforts align with broader (CWR) strategies, emphasizing systematic evaluation and integration to enhance resilience in breeding programs. Challenges in genetic resource management include collection gaps, particularly in southern where lower and higher homozygosity limit representation. Post-2020 genome sequencing has highlighted the need for updated assessments to capture structural variants and refine core collections. International initiatives follow FAO guidelines for , promoting standardized operations and global exchange to support and utilization. These frameworks ensure S. pimpinellifolium contributes to amid climate pressures.

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