diff --git a/src/View/Antlers/Parser.php b/src/View/Antlers/Parser.php
index 3017bd05c0..f230d306b2 100644
--- a/src/View/Antlers/Parser.php
+++ b/src/View/Antlers/Parser.php
@@ -66,10 +66,10 @@ class Parser
     public function __construct()
     {
         // Matches a variable inside curly braces. Spaces not allowed.
-        $this->variableRegex = "(?!if\s|unless\s)[a-zA-Z0-9_'\"][^<>{}=\s]*";
+        $this->variableRegex = "(?!if\s|unless\s)[a-zA-Z0-9_'\"][^{}=\s]*";
 
         // Matches a full variable expression inside curly braces.
-        $this->looseVariableRegex = "(?!if\s|unless\s)[a-zA-Z0-9_'\"][^<>{}=]*";
+        $this->looseVariableRegex = "(?!if\s|unless\s)[a-zA-Z0-9_'\"][^{}=]*";
 
         // Matches the first part of a {{ tag: followed a variable name and full expression.
         $this->callbackNameRegex = '(?!if\s|unless\s)[a-zA-Z0-9_][^<>{}=!?]*'.':'.$this->variableRegex;
diff --git a/tests/View/Antlers/ParserTest.php b/tests/View/Antlers/ParserTest.php
index 1e67566bcd..0277613ec7 100644
--- a/tests/View/Antlers/ParserTest.php
+++ b/tests/View/Antlers/ParserTest.php
@@ -413,6 +413,16 @@ public function testNullCoalescence()
         $this->assertEquals('Pass', $this->parse('{{ missing[thing] ?? "Pass" }}', $this->variables));
     }
 
+    public function testNullCoalescenceWithStringModifiers()
+    {
+        $this->assertEquals('HELLO WILDERNESS', $this->parse('{{ missing ?? string | upper }}', $this->variables));
+        $this->assertEquals('HELLO WILDERNESS', $this->parse('{{ missing | upper ?? string | upper }}', $this->variables));
+        $this->assertEquals('HELLO WILDERNESS', $this->parse('{{ string | upper ?? missing | upper }}', $this->variables));
+
+        // With parameters
+        $this->assertEquals('Hello wilderness >', $this->parse('{{ missing ?? string | ensure_right: > }}', $this->variables));
+    }
+
     public function testTruthCoalescing()
     {
         $this->assertEquals('Pass', $this->parse('{{ string ?= "Pass" }}', $this->variables));