Pinaceae

Pinaceae Temporal range: Jurassic–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Larix (golden), Abies (central foreground) and Pinus (right foreground)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Gymnospermae
Division:	Pinophyta
Class:	Pinopsida
Order:	Pinales
Family:	Pinaceae Lindley 1836
Genera
Abietoideae Abies Cedrus Keteleeria Nothotsuga Pseudolarix Tsuga Pinoideae Pinus Piceoideae Picea Laricoideae Pseudotsuga Cathaya Larix
Synonyms
Abietaceae von Berchtold & Presl 1820 Cedraceae Vest 1818 Compsostrobaceae Delevoryas & Hope 1973 †Kranneraceae Corda 1866 Piceaceae Goroschankin 1904

The Pinaceae (/pɪˈneɪsiːˌiː, -siˌaɪ/), or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as cedars, firs, hemlocks, piñons,

Members of the family Pinaceae are

The female cones are large and usually woody, 2–60 centimetres (1–24 inches) long, with numerous spirally arranged scales, and two winged seeds on each scale. The male cones are small, 0.5–6 cm (1⁄4–2+1⁄4 in) long, and fall soon after pollination; pollen dispersal is by wind. Seed dispersal is mostly by wind, but some species have large seeds with reduced wings, and are dispersed by birds. Analysis of Pinaceae cones reveals how selective pressure has shaped the evolution of variable cone size and function throughout the family. Variation in cone size in the family has likely resulted from the variation of seed dispersal mechanisms available in their environments over time. All Pinaceae with seeds weighing less than 90 milligrams are seemingly adapted for wind dispersal. Pines having seeds larger than 100 mg are more likely to have benefited from adaptations that promote animal dispersal, particularly by birds.^[4] Pinaceae that persist in areas where tree squirrels are abundant do not seem to have evolved adaptations for bird dispersal.

Boreal conifers have many adaptions for winter. The narrow conical shape of northern conifers, and their downward-drooping limbs help them shed snow, and many of them seasonally alter their biochemistry to make them more resistant to freezing, called "hardening".

Classification

Classification of the subfamilies and genera of Pinaceae has been subject to debate in the past. Pinaceae ecology, morphology, and history have all been used as the basis for methods of analyses of the family. An 1891 publication divided the family into two subfamilies, using the number and position of resin canals in the primary vascular region of the young taproot as the primary consideration. In a 1910 publication, the family was divided into two tribes based on the occurrence and type of long–short shoot dimorphism.

A more recent classification divided the subfamilies and genera based on the consideration of features of ovulate cone anatomy among extant and fossil members of the family. Below is an example of how the morphology has been used to classify Pinaceae. The 11 genera are grouped into four subfamilies, based on the microscopical anatomy and the morphology of the cones, pollen, wood, seeds, and leaves:^[5]

• Subfamily
  Pinus): cones are biennial, rarely triennial, with each year's scale-growth distinct, forming an umbo on each scale, the cone scale base is broad, concealing the seeds fully from abaxial (below the phloem
  vessels) view, the seed is without resin vesicles, the seed wing holds the seed in a pair of claws, leaves have primary stomatal bands adaxial (above the xylem) or equally on both surfaces.
• Subfamily
  Picea
  ): cones are annual, without a distinct umbo, the cone scale base is broad, concealing the seeds fully from abaxial view, seed is without resin vesicles, blackish, the seed wing holds the seed loosely in a cup, leaves have primary stomatal bands adaxial (above the xylem) or equally on both surfaces.
• Subfamily
  Larix, Pseudotsuga, and Cathaya
  ): cones are annual, without a distinct umbo, the cone scale base is broad, concealing the seeds fully from abaxial view, the seed is without resin vesicles, whitish, the seed wing holds the seed tightly in a cup, leaves have primary stomatal bands abaxial only.
• Subfamily
  Abies, Cedrus, Pseudolarix, Keteleeria, Nothotsuga, and Tsuga
  ): cones are annual, without a distinct umbo, the cone scale base is narrow, with the seeds partly visible in abaxial view, the seed has resin vesicles, the seed wing holds the seed tightly in a cup, leaves have primary stomatal bands abaxial only.

Multiple molecular studies indicate that in contrast to previous classifications placing it outside the conifers, Gnetophyta may in fact be the sister group to the Pinaceae, with both lineages having diverged during the early-mid Carboniferous. This is known as the "gnepine" hypothesis.^[11][12]

Evolutionary history

Pinaceae is estimated to have diverged from other conifer groups during the late

External stresses on plants have the ability to change the structure and composition of forest ecosystems. Common external stress that Pinaceae experience are herbivore and pathogen attack which often leads to tree death.^[22] In order to combat these stresses, trees need to adapt or evolve defenses against these stresses. Pinaceae have evolved a myriad of mechanical and chemical defenses, or a combination of the two, in order to protect themselves against antagonists.^[23] Pinaceae have the ability to up-regulate a combination of constitutive mechanical and chemical strategies to further their defenses.^[24]

Pinaceae defenses are prevalent in the bark of the trees. This part of the tree contributes a complex defensive boundary against external antagonists.^[25] Constitutive and induced defenses are both found in the bark.^[25][26][27]

Constitutive defenses

Constitutive defenses are typically the first line of defenses used against antagonists and can include sclerified cells, lignified periderm cells, and secondary compounds such as phenolics and resins.^[28][25][26] Constitutive defenses are always expressed and offer immediate protection from invaders but could also be defeated by antagonists that have evolved adaptations to these defense mechanisms.^[28][25] One of the common secondary compounds used by Pinaceae are phenolics or polyphenols. These secondary compounds are preserved in vacuoles of polyphenolic parenchyma cells (PP) in the secondary phloem.^[29][27]

Induced defenses

Induced defense responses need to be activated by certain cues, such as herbivore damage or other biotic signals.^[28]

A common induced defense mechanism used by Pinaceae is resins.

The topic of defense mechanisms within family Pinaceae is a very active area of study with numerous studies being conducted. Many of these studies use methyl jasmonate (MJ) as an antagonist.^[26][27][32] Methyl jasmonate is known to be able to induce defense responses in the stems of multiple Pinaceae species.^[26][32] It has been found that MJ stimulated the activation of PP cells and formation of xylem traumatic resin ducts (TD). These are structures that are involved in the release of phenolics and resins, both forms of defense mechanism.^[26][27]

• 
  Close up of bishop pine cones
• 
  Knobcone pine cone

References

External links

Wikimedia Commons has media related to Pinaceae.

Wikispecies has information related to Pinaceae.

• Arboretum de Villardebelle French Arboretum of conifers around the world
• Gymnosperm Database – Pinaceae
• Pinaceae on the web page of the Tree-of-Life project
• 40 Pine Trees From Around the World by The Spruce
• Jepson eFlora, The Jepson Herbarium, University of California, Berkeley, covers Californian species and much of western North America
• Pinaceae in Flora of North America
• Pinus in USDA Plants Database